Pregnant. by Contrast, a Pregnancy Rate of 88 Percent Was Evident

AN ABSTRACT OF THE THESIS OF

CHARLES EDWARD TRAINER for the MASTER OF SCIENCE (Name) (Degree) in WILDLIFE MANAGEMENT presented on (Major) ci(Date) Title: THE RELATIONSHIP OF PHYSICAL CONDITION AND FERTILITY OF FEMALE ROOSEVELT ELK (CERVUS

CANADENSIS ROOSEVELTI) IN OREGON Redacted for Privacy Abstract approved: Howard M. Wight

The ratio of calves to cows observed annually during winters of 1950-69 for Roosevelt elk in Western Oregon indicated that about ZO percent fewer calves were seen than in similar inventories for Rocky Mountain elk (Cervus canadensis nelsorli) in Northeast Oregon. To determine if fertility was an important cause of the comparatively low percentage of calves recorded for Roosevelt elk, factors affecting reproduction in the subspecies were investigated. Examination of Z64 uteri of adult (two years and older) Roosevelt elk collected on three study areas in Western Oregon during

the November-January periods,1964-68revealed that 50 percent were pregnant. By contrast, a pregnancy rate of 88 percent was evident for 90 adult Rocky Mountain elk harvested in Northeast Oregon between November and January,1967-68and1968-69.Comparison of the average 50 percent rate of pregnancy apparent for adult Roosevelt elk examined during 1964-68 with the mean calf: cow ratio of 41 percent observed about 13 months later, indicates that the proportion of calves enumerated was largely determined by fertility rather than postnatal survival.Different percentages of pregnant cows were evident among various age groups of Roosevelt elk; yearlings (12per- cent), two and three year olds (32 percent), four to ten year olds (59 percent) and all of the eight cows examined, estimated to be older than ten years, were nongravid. None of 14 nonpregnant yearling Roosevelt elk sampled had ovulated, and the evidence of infertility identified in a sample of 79 nongravid adults intensively studied was apparent as embryonic mortality (minimum of two percent) and anovulation (23 to 64 percent). Tests for diseases harmful to reproduction indicated that pathological factors were not affecting fertility.An indicated decline in the ratio of bulls in Roosevelt elk herds (a result of increased harvests) coupled with no apparent decrease in calf: cow ratios suggested that the relatively low fertility rate in the subspecies originated with the female. Measurements of kidney fat depots indicated significant differences in physical condition between pregnant and nonpregnant cows that were not revealed by adrenal weight indices. Among cows 3-10 years old, there was evidence of significantly larger kidney fat deposits in gravid than in nongravid females.Lactating cows (3-10 years old) were indicated to have significantly smaller depots of kidney fat and a lower pregnancy rate (51 percent pregnant) than nonlactating cows (84 percent pregnant).Earlier dates of breeding were generally apparent for adult cows in the best physical condition. The relationship between fat stores and fertility evident for female Roosevelt elk indicates that environmental factors, probably re- 1.ated to nutrition, were the underlying cause of the low reproductive rates observed in the subspecies.The stress of lactation was identified as the main physiological process affecting energy reserves of the cow to the detriment of fertility. Data were not available to determine if reproduction was limited because of qualitative or quantitative deficiencies in forage.Research concerning the effect of various population densities of Roosevelt elk upon fertility is suggested to determine whether or not the present level of reproduction represents the maximum net productivity possible under prevailing environmental conditions. The Relationship of Physical Condition and Fertility of Female Roosevelt Elk (Cervus canadensis roosevelti) in Oregon

by Charles Edward Trainer

A THESIS submitted to Oregon State University

in partial fulfillment of the requirements for the degree of Master of Science June 1971 APPROVED:

Redacted for Privacy

Professor of Wildlife Ecology in charge of major

Redacted for Privacy

Head of Department of Fisheries and Wildlife

Redacted for Privacy

Dean of Graduate School

Date thesis is presented Typed by Donna L. Olson for Charles Edward Trainer ACKNOWLEDGEMENTS

My sincere appreciation is expressed to Mr. Howard Wight, Professor of Wildlife Ecology, Department of Fisheries and Wildlife, for his careful guidance during the study.I am also grateful to the Oregon State Game Commission for supporting the investigation under Pittman-Robertson Project W-59-R. This study would not have been possible without the cooperation of many individuals.I wish particularly to thank Mr. James Harper and Mr. William Lightfoot for many long and often difficult days spent in the planning and supervision of special hunting seasons involved with collection of elk.Special recognition is due to Messrs. Lyle Carver, Kenneth Cochrun, William Hall, Francis Ives, Frank LeMay, Robert Mace, John Rayner, Robert Stein, Harold Sturgis and Donald Wilt for their assistance with the intensive collection of specimens on the Millicoma area.I also wish to thank other Game Commission personnel, members of the State Police and hunters who submitted much material for examination and aided the study in other ways. Appreciation is extended to Mr. Warren Aney and Dr. W. Scott Overton for their aid with statistical analyses, and to Dr. Dean Smith for conducting tests for pathogens.I am grateful to DrPaul Vohs for his thoughtful criticism of the manuscript. Dr. Kenneth Greer, Montana State Department of Fish and Game, Dr. B. W. O'Gara, U. S. Fish and Wildlife Service and Dr. Philip Wright, University of Montana, provided many helpful sugges- tions concerning the laboratory work.Their assistance is gratefully appreciated. Above all I wish to acknowledge my wife, Clara, and daughters Audrey and Kimberly for their willing acceptance of the many hours which I spent apart from the family to complete the research and thesis. TABLE OF CONTENTS

Chapter Page

I.INTRODUCTION 1

II. METHODS 4

Collection of Specimens 4 Laboratory Methods 6 Terminology 9

III. RESULTS 11

Breeding Season 11 Breeding Seasons of 1967 and 1968 11 Physical Condition and Period of Breeding 15 Fertility 20 Pregnancy Rates 20 Incidences of Twinning and Multiple Ovulation 26 Evidence of Infertility 27 Infertility inYearlings 27 Infertility in Adults 28 Anovulation 28 Embryonic Mortality 30 Infertility in Rocky Mountain Elk 32 Factors Influencing Fertility 34 Pathology 34 Diseases Associated with Reproduction 34 Abnormalities in Uteri and Ovaries 35 Male Coverage 36 Physical Condition 37 Indices of Physical Condition 37 Indices of Kidney and Marrow Fat 40 Indices of Adrenal Weight 48

IV.DISCUSSION 53

V. CONCLUSIONS 62

LITERATURE CITED 64

APPENDICES 68 LIST OF FIGURES

Figure Page

1. Present distribution of elk in Oregon and study areas where Roosevelt elk were collected, 1964- 68 reproductive seasons. 5 2. Relationship of kidney fat indices and dates of breeding for 25 female Roosevelt elk, 3-10 years old, collected in Western Oregon, November 2-17, 1968. 17 3. Relationship of kidney fat indices and dates of breeding for 20 female Roosevelt elk, 3-10 years old, collected in Western Oregon, January 6-28, 1968. 18 4. Kidney fat indices plotted against percentage of fat (wet basis) in metatarsal marrow for 54 female Roosevelt elk, three years and older, collected in Western Oregon, November 2-December 15, 1968. 41 5. Kidney fat indices plotted against percentage of fat (wet basis) in metatarsal marrow for 41 female Roosevelt elk, three years and older, collected in Western Oregon, January 3-28, 1968. 42 LIST OF TABLES

Table Page

1. Dates of breeding calculated for adult female Roosevelt elk collected in Western Oregon, 1967 and 1968 reproductive seasons (dates of collection in parenthesis). 12 2. Dates of breeding calculated for adult female Rocky Mountain elk collected in Northeast Oregon, 1967 and 1968 reproductive seasons (dates of collection in parenthesis). 13 3. Pregnancy rates of Roosevelt elk collected in Western Oregon, 1964-68 reproductive seasons (dates of collection in parenthesis). 21 4. Comparison of age specific pregnancy rates among samples of Roosevelt and Rocky Mountain elk (outside collection dates in parenthesis--all years listed as reproductive seasons), 25

5. Infertility identified in nongravid adult Roosevelt elk collected in Western Oregon, 1965, 1967 and 1968 reproductive seasons. 29

6. Description of occurrences and suspected occurrences of embryonic mortality for Roosevelt elk collected in Western Oregon, 1965, 1967 and 1968 reproductive seasons. 31

7, Infertility identified in nongravid Rocky Mountain elk collected in Northeast Oregon, 1967 and 1968 reproductive seasons. 33

8. Ratios of bulls and calves per 100 cows recorded annually for Roosevelt elk in Western Oregon between December and mid-February, 1950-70,

9. Indices of kidney fat for yearling and two year old female Roosevelt elk collected in Western Oregon, 1968 reproductive season; all specimens nonlactating. 44 Table Page

10. Relationship of pregnancy rates and status of lactation for Roosevelt elk, 3-10 years old, collected in Western Oregon, 1967 and 1968 reproductive seasons. 46 11. Relationship of pregnancy rates and status of lactation for Rocky Mountain elk, 3-10 years old, collected in Northeast Oregon, 1967 and 1968 reproductive seasons. 47

1Z. Correlation coefficients (r) computed from correlation of three indices of adrenal weight with variables listed for female Roosevelt elk, 3-10 years old, collected on Millicoma area, 1967 and 1968 reproductive seasons, 50

13. Coefficients of determination (R2) and F values resulting from stepwise multiple linear regression analysis of adrenal weight: hog-dressed body weight ratio Y on variables listed for female Roosevelt elk, 3-10 years old, collected on Millicoma area, 1967 and 1968 reproductive seasons. 51 LIST OF APPENDIX TABLES

Appendix 1 Page Table A. Ovarian and uterine data--pregnant Roosevelt elk collected on Millicoma area, November 20 and 21, 1965. 69 B. Ovarian and uterine data- -pregnant Roosevelt elk collected on Millicoma area, January 6-28, 1968. 70

C. Ovarian and uterine data--pregnant Roosevelt elk collected on Millicoma area, November 2 - December 14, 1968. 71 Ovarian and uterine data--pregnant Roosevelt elk collected on North Coast area, November 11-18, 1967. 72 E. Ovarian and uterine data--pregnant Roosevelt elk collected on North Coast area, November 16 and 17, 1968 and February 15, 1969. 73 F. Ovarian and uterine data--pregnant Roosevelt elk collected on Loon Lake area, November 17 - December 15, 1968. 74

G. Ovarian and uterine data- -nonpregnant Roosevelt elk collected on Millicoma area, November 20 and 21, 1965. 75

1-1. Ovarian and uterine data--nonpregnant Roosevelt elk collected on Millicoma area, January 3February 25, 1968. 76 I. Ovarian and uterine data--nonpregnant Roosevelt elk collected on Millicoma area, November 2 - December 14, 1968. 77 J. Ovarian and uterine data--nonpregnant Roosevelt elk collected on North Coast area, November 11 - December 6, 1967. 78 Appendix 1 Table Page

K. Ovarian and uterine data- -nonpregnant Roosevelt elk collected on North Coast area, November 16 and 17, 1968, 79 L. Ovarian and uterine data- -nonpregnant Roosevelt elk collected on Loon Lake area, December 7-15, 1968. 80

Appendix Z Table

A. Ovarian and uterine data- -pregnant Rocky Mountain elk collected in Northeast Oregon, November 11, 1967 - January ZO, 1968. 8Z B. Ovarian and uterine data--pregnant Rocky Mountain elk collected in Northeast Oregon, November 16, 1968 - January Z5, 1969. 83 C. Ovarian and uterine data- -nonpregnant Rocky Mountain elk collected in Northeast Oregon, November 11-19, 1967. 85

D. Ovarian and uterine data- -nonpregnant Rocky Mountain elk collected in Northeast Oregon, November 16 and 17, and December 22, 1968. 86 Appendix Table 3. Sex of Roosevelt and Rocky Mountain elk fetuses collected in Oregon, 1965, 1967 and 1968 reproductive seasons, 87 4. Data on physical condition for pregnant Roosevelt elk collected on Millicoma area, 1967 reproductive season, 88

5. Data on physical condition for nonpregnant Roosevelt elk collected on Millicoma area, 1967 reproductive season, 89 Appendix Table Page

6, Data on physical condition for pregnant Roosevelt elk collected in Western Oregon, 1968reproductive season, 90

7. Data on physical condition for nonpregnant Roosevelt elk collected in Western Oregon, 1968reproductive season. 9Z THE RELATIONSHIP OF PHYSICAL CONDITION AND FERTILITY OF FEMALE ROOSEVELT ELK (CERVUS CANADENSIS ROOSEVELTI) IN OREGON

I.INTRODUCTION

The purpose of this study was to relate physical condition and other factors that influence reproduction to fertility rates observed for female Roosevelt elk in Oregon.It is hoped that data presented will increase understanding of factors affecting reproduction in Roosevelt elk populations and thereby contribute to knowledge essen- tial for management of the herds. Two subspecies of elk occur in Oregon. Mace (1956) reports that Roosevelt elk are distributed intermittently throughout the Coast and Cascade Mountain ranges, whereas Rocky Mountain elk are found in the northeast section of the state, principally in the Blue and Wallowa mountains (Figure 1). Schwartz and Mitchell (1945: 311) recorded an average of 61 calves per 100 cows for 419 Roosevelt elk cows and Z55 calves classified on the Olympic Peninsula, Washington in July and August of 1936- 38 But ratios of calves to cows on Afognak Island, Alaska averaged only 3Z: 100 during the summers of 1961 -63 (Batchelor, 1965: 14) Calf: cow ratios obtained for Roosevelt elk in Oregon from December a

to mid-February, 1950-69 revealed an average of 41 calvesper 100 cows for 11,522 calves and 27,938 cows classified.Similar data for Rocky Mountain elk, involving observations of 20, 013 calves and 39. 538 cows during the same period in Northeast Oregon averaged 51 calves per 100 cows or 20 percent more calves than for Roosevelt elk. The ratio of calves to cows indicated for Roosevelt elk suggested that many cows might not produce an offspring,or their young died, or cows were more readily observable than calves.Interpreting counts of calves, therefore, required knowledge of the proportion of femaLes likely to be pregnant.To fill this need, uteri from Roosevelt elk killed during 1964, 1965 and 1966 on the Millicoma Tree Farm in Southwest Oregon were examined for evidence ofpregnancy. Of 125 females (two years and older) examined, only 46 percentwere gravid

(Harper, 1965:7; unpublished field notes, 1965; and 1967: 3).By comparison, reproductive rates calculated from uterine analysis of adult (two years and older) Rocky Mountain elk were much higher. Kittams (1953: 183) reported a pregnancy rate of 85 percent for 1,053 adult cows killed in Yellowstone National Park between 1935 and 1951. Greer (1966: 124) found that from 80 to 95 percent of the adult females comprising samples collected annually in Yellowstone Park from 1961 to 1965 were gravid. Consideration of the markedly different reproductive rates 3 observed for the two subspecies raised the question of why rates were lower in Roosevelt elk.Consequently, my study was initiated in Noember, 1967 to investigate certain factors affecting fertility in Roosevelt elk; specifically to:

(1) determine the fertility rate.

(2) identify, insofar as possible, factors that inhibit fertility.

(3) determj.ne the duration of the breeding season.

(4) assess the physical condition of cows examined to relate condition of the animal to fertility. 4 II. METHODS

Collection of Specimens

The present study is based on examination of 289 uteri of Roosevelt elk killed in the Coast Range of Oregon from 1964 to 1968. No specimens were obtained earlier than November or later than February each year. James Harper analyzed uteri collected prior to

1967. The specimens originated from three locations that I have designated as North Coast, Loon Lake, and Millicoma study areas (Figure

1).Although high densities of elk prevailed in the three localities, most specimens (240) were from the Millicoma area- -a 110-square mile portion of the Millicoma Tree Farm lying north and northeast of Coos Ridge. To compare with similar data on Roosevelt elk, biologists collected uteri from 105 Rocky Mountain elk harvested throughout North- east Oregon during the November-January periods, 1967 and 1968, Three methods were employed to gather materials for study. Hunters participating in antlerless -only elk seasons were briefed about the collecting program and asked to save reproductive tracts, udders, and mandibles from cows they killed.Appropriate instructions, dia- grams, and storage bags were supplied.The completeness of samples received from hunters seemed related to the amount of time spent 5

0000000 000000 0 / 0 0 00 00 0 0 00 o0 0 7/ 00 o'00°0 b/ 0 0 0000 //// 000 000000 0 0 0 0 0 0 00 / 00000 00 -12) / /

Legend

Roosevelt Elk Distribution

0 0 00 0000 = Rocky Mountain Elk Distribution - - - - =Study Area Boundary

Study Areas: 1.= North Coast 3 = Millicoma 2 = Loon Lake

Figure 1.Present distribution of elk in Oregon (Mace, 1956) and study areas where Roosevelt elk were collected, 1964-68 reproductive seasons. instructing the individual. Personnel concerned with enforcement duties also submitted materials from cows killed illegally. Supervised hunts were conducted during January, 1968 and November, 1968 on the Millicoma area to provide Rooseveltelk for intensive study. From six to ten huntersper day were scheduled during each hunt. On the appointed days, biologistsaccompanied the parties afield and obtained the following items from eachelk killed: whole and hog-dressed weights, the reproductivetract, mandible, udder, kidney fat depots, adrenals,one metatarsal bone, and a sample of blood.Both hunts proved successful from standpoints of datacol- lected and satisfaction of hunters.

Uteri were preserved in AFA (Mossman's recipe--Guyer,1953: 236) or by freezing.Other materials were kept coolor frozen until examined.

Laboratory Methods

Study of reproductive tracts under laboratory conditionsincluded examination of the ovaries, uterine contents, cervix andvagina.Tis - sues for histological study were sectioned at seven microns and stained with Harris' hematoxylin and Putt's eosin,

Ovaries (fixed) were sliced along the longitudinal axis witha razor blade into sections 1 to 2. mm thick, leaving the sections 7 attached to the mesovarium. The average diameter (measured at right angles) of corpora lutea, corpora albicantia, and all follicles larger than 2 mm was then tabulated. All uteri were dissected and examined for evidence of pregnancy.Uteri showing an initial enlargement typical of early pregnancy, or with ovaries containing corpora lutea, were severed from the tract at the mid-cervical region and trimmed of mesometrial tissue.The uterine horns were then opened while submerged in tap water, and a search was conducted for small, macroscopic embryonic tissue.No attempt was made to recover ova or microscopic-sized blastocysts.The cervical and vaginal mucosa were also examined for abnormalities. Embryos were removed from gravid uteri, scrutinized for ex- ternal anomalies and the sex, body length, and weight recorded. Following procedures outlined by Armstrong (1950: 652) and Morrison, Trainer and Wright (1959: 29), I used two methods of measuring length in embryos. These were: Crown-rump length -- For specimens up to 64 mm, the distance between the anterior-most to the posterior-most points of the body was measured. Forehead-rump length -- For specimens larger than 64mm, the embryo was placed on its side with its back againsta straight surface and care was taken to prevent the head from curling tailward.The distance measured was from the anterior-most point of the crown to the tuberosity of the ischium. Dates of conception were calculated usinga growth curve de- rived from Rocky Mountain elk embryos of knownage (Morrison, Trainer and Wright, 1959).

Ages of elk were estimated at the birthdate previousto date of collection according to tooth replacement andwear patterns described by Quimby and Gaab (1957). With one modification, deposits of kidney fatwere measured using the kidney fat index (weight of fat surrounding the kidneys divided by weight of the kidneys) described by Riney (1955: 433and 434).Riney cut away and discarded fat anterior and posteriorto the kidney before weighing the remaining fat.To avoid variation in main- taming a900cutting plane when trimming fat from ends of the kidneys (B. W. OGara, personal communication, 1967), calculationof the index in my study (based on both kidneys and expressed in percent)included all fat attached to the kidneys.Therefore the kidney fat indices presented are larger than would have been thecase using Riney's technique.

Fat content of marrow samples from the metatarsalswas determined on a wet and dry weight basis by the Department ofAnimal Science, Oregon State University, following standardprocedures for ether extraction.The marrows (in situ) were frozen from 45 to 60 days before analysis. The adrenal glands were dissected from the kidneys, fixed in ten percent formalin, and stored in 70 percent alcohol.After fat was removed from preserved adrenals, the glands were blotted dry and weighed to the nearest 0. 01 gram. Samples of blood serum were tested for Brucella abortus, Leptospira pomona, L. canicola, L. icterohaemorrhagiae, and Anaplasrna marginale by the Veterinary Diagnostic Laboratory, Oregon State University.The tests employed were the serum plate agglutination test of the United States Department of Agriculture, the rapid plate agglutination test, and theanatest',respectively. The diagnostic laboratory also checked samples of cervical mucus for Vibrio fetus (var. veneralis) using culturing, fluorescent antibody, and cervical mucus agglutination techniques.

Terminology

Pregnancy rate is used to express the percentage of pregnant females in a sample consisting of adult females (two years and older) or yearling females. Since elk rarely bear more than one calf (Kittams, 1953: 18Z and 183), fertility and fecundity are used synony- mously with pregnancy rate.

"DiamondLaboratories, Des Moines, Iowa. Reproductive season refers to the year in which the rut occurred.This designation was necessary because collections from cne gestational period often extended over part of two calendar years. Embryo is used in a broad sense to include specimens in all phases of prenatal development. Statistical significance was considered to be at the 0, 05 level unless otherwise stated. 11 III. RESULTS

Breeding Season

Breeding Seasons of 1967 and 1968

Murie (1951: 125) reported that the active breedingseason for elk occurs from the first part of September to the latterpart of October.His observations were based on intensity of bugling and other rutting behavior of bulls.Judging from dates calves were dropped, Schwartz and Mitchell (1945:298) indicated that most breeding of Roosevelt elk on the Olympic Peninsula, Washington, took place between September 15 and October 15,Based on dates of conception computed from embryo size, Morrison, Trainer and Wright (1959: 31) found that 26 (65 percent) of 40 gravidcows collected from two Rocky Mountain elk populations in Montana conceived between September 26 and October 10, whereas outside dates of breedingwere September 16, and November 4. Dates of conception were calculated for Roosevelt and Rocky Mountain elk embryos collected during the 1967 and 1968 reproductive seasons to delimit the period of the rut,Comparison of respective distributions of breeding dates for adultcows of both subspecies mdi- cates that females collected in November generally conceived earlier than did cows obtained during December and January (Tables 1and 2), Table 1.Dates of breeding calculated for adult female Roosevelt elk collectedin Western Oregon, 1967 and 1968 reproductive seasons (dates of collection in parenthesis).

1967 Season 1968 Season 1967&1968 Seasons November January November December November Post November (11-18) (6-28, 1968) (2-22) (7-15) Cumulative Cumulative Dates of Breeding No. No. No. No. No. Percent No. Percent Aug. 17-21 1 1 2.7 22-26 27-31 Sept. 1 1- 5 2 3 10.8 6-10 5 5 24.3 11-15 1 1 5 6 40.5 1 2.8 16-20 1 2 3 4 51.3 2 8.3 21-25 2 1 4 1 54.1 6 25.0 26-30 1 4 4 3 5 67.6 7 44.4 Oct. 1- 5 6 5 1 5 81.1 7 63.9 6-10 4 1 1 83.8 4 75.0 11-15 1 3 3 91.9 1 77.8 16-20 2 1 1 1 94.6 3 86.1 21-25 1 1 1 97.3 1 88.9 26-30 1 1 1 1 1000 2 94.4 31- 4 Nov. 1 1 97.2 Nov. 5- 9 97.2 10-14 97.2 15-19 1 1 100.0 Totals 5 24 32 12 37 36 Earliest Date Sept.5 Sept.15 Aug. 19 22 " Sept. Aug.19 Sept. 15 Latest Oct.22 Nov. 3 Oct.30 Nov.17 Oct.30 Nov. 17 Median " Sept. 20 Oct. 3 Sept. 18 Sept.28 Sept.20 Oct. 2 Mean " Sept. 24 Oct. 4 Sept. 23 Oct. 7 Sept.23 Oct. 5

I- (") Table 2.Dates of breeding calculated for adult female Rocky Mountain elk collected in Northeast Oregon, 1967 and 1968 reproductive seasons (dates of collection in parenthesis).

1967 Season 1968 Season 1967 G 1968 Seasons November Dec.11, 67 November Dec.7, 68 November Post November (11-19) Jan.20, 68 (16-20) Jan.25, 69 Cumulative Cumulative No. No. No. No. Dates of Breeding No. Percent No. Percent

Sept. 6-10 2 2 3.5 11-15 2 1 2 7.0 1 5.6 16-20 1 5 1 6 17.5 1 11.1 21-25 1 4 3 5 26.3 3 27.8 26-30 9 1 2 2 11 45.6 3 44.4 Oct. 1- 5 9 2 6 1 15 71.9 3 61.1 6-10 6 4 2 10 89.5 2 72.0 11-15 2 1 2 93.0 1 77.8 16-20 2 1 2 96.5 1 83.3 21-25 1 1 1 98.2 1 88.9 26-30 1 1 100.0 88.9 31- 4 Nov. 1 1 94.4 Nov. 5- 9 1 1 100.0

Totals 29 6 28 12 57 18

EarliestDate Sept. 16 Sept.27 Sept. 10 Sept.13 Sept.10 Sept.13 Latest " Oct.24 Nov. 8 Oct.27 Oct. 17 Oct. 27 Nov. 8 Median Oct. 3 Oct. 5 Sept. 28 Sept.28 Oct. 2 Oct. 2 Mean " Oct. 3 Oct. 17 Sept. 28 Sept.30 Oct. 1 Oct. 1 14 The differences observed could not beattributed to any variation in time of breeding activity betweenareas where elk were collected Although some of the difference in chronologiesof breeding dates between the November and post November periodsmight be explained by random variation in the small samples,at least one other factor is apparently also involved.

Inspection of the growth curve for knownage elk embryos (Morrison, Trainer and Wright, 1959)suggests that approximately 20 days postconception is the minimum timeneeded for embryonic tissue in elk to become macroscopic, anddetectable with the techniques applied in my study.Since uteri of adult cows were collected beginning November 2, I believe that the consistentlyearlier cessation of breeding indicated for November collectionsresulted primarily from a failure to identify all conceptions occurring after October 13 (20 days prior to November 2).For the 1967 and 1968 reproductive seasons combined, distribution of breeding dates determined forcows collected during the December-January periodsuggests that six percent (two of 36) and 11 percent (two of 18) of the Rooseveltand Rocky Mountain elk (adults), respectively conceivedafter the last breeding dates indicated by the November collections(Tables 1 and 2)Conse- quently,the breeding dates designated forcows obtained in November probably accurately represent the chronology ofrutting activity to only about 20 days before the first daycows were collected (October 15 13).The distributions of conceptions calculated for the December- January samples appear then to most accurately describe the breeding season for el.k examined.

The breeding seasons determined for adult Roosevelt andRocky Mountain elk during the 1967 and 1968 reproductiveseasons (December- January samples) are similar in most respects. Also the dates of breeding agree well with the timing of the rut in both subspeciesas re-

ported by Murie (1951; 125), Schwartz and Mitchell (1945:298)arid Morrison, Trainer and Wright (1959: 31). Although the dates that adult cowswere bred evidently was not related to their age, yearlings usually conceived later than didadults. For Rocky Mountain elk, the average breeding date for five yearlings collected during November, 1968 was October 6 (range, September

29 - October 10) compared to September 28 for adults obtainedin November (Table 2).Two gravid yearling Roosevelt elk killed December 7, 1968 and February 15, 1969 were estimatedto have conceived on October 13 and November 10, respectively, whereasthe mean date of breeding for adults obtained in December, 1968was October 7 (Table 1).

Physical Condition and Period of Breeding

Verme (1965) reported that the breeding season of adult female white-tailed deer (Odocoileus virginianus) subjectedto extremely poor 16

nutrition from October to late November was later than for doeson a high dietary plane during the same period.The rutting season of fe-

male Tule elk (Cervus canadensis nannodes) that McCulloch(1969: 59

and60)studied occurred earlier during years of abundant forageas compared to years of poor food production.Apparently the nutritional status of female Roosevelt elk during the fall (as judged by their physical condition) influenced dates that cows conceived. For a sample of 25 cows, 3-10 years old, collected during

November,1968,females judged in the best physical condition (mdi-

cated by higher kidney fat indices) had earlier breeding dateson the average than did cows in poorer condition (Figure 2).The relationship between improved physical condition and earlier breedingwas weaker, however, for a collection of 20 cows (3-10 years old)

examined during January,1968(Figure 3).But the interval between time of collection and peak of the respective1967and1968ruts was about90days for cows obtained in January,1968compared to approximately 30 days for specimens examined in November,1968(Table 1).

Consequently, had females collected during January,1968been examined in November,1967(thus considerably reducing the effect of interim environmental factors upon fat deposits) then possiblya strong- er relationship between physical condition and time of breeding would have been apparent for cows obtained in January.

Lactating Roosevelt elk, presumably because of generallypoorer Indices of Kidney Fat (Classes) 195_284[ 9/10

165 - 194

9/18 = Breeding Date (Dry Cow) = Breeding Date (Wet Cow) 0 - = Mean Breeding Date 101- 164. B I 9/24

I 63- 100 Ai I 9/24

32- 62I- A I 9/29

1968 30 10 20 30 10 20 30 August September October Breeding Dates

Figure 2. Relationsliip of kidney fat indices and datesof breeding for 25 female Roosevelt elk, 3-10years old, collected inWestern Oregon, November 2-17, 1968 Indices of Kidney Fat (Classes)

81 - 127 A 9/30

I ° U 61-80 I' 10/10 = Breeding Date (Dry Cow) C) = Breeding Date (Wet Cow) 43- 57 A = Mean Breeding Date 9/28

5) c-. C) - 14- 33 A 10/7

I I I I 1967 10 20 30 10 20 30 10 September October November Breeding Dates

Figure 3..Relationship of kidney fat indices and dates of breeding for 20 female Roosevelt elk, 3-10years old, collected in Western Oregon, January 6-28, 1968. 19 physical condition, usually becamepregnant later than did nonlactating females (Figures 2 and 3).Fuller (1966: 21) reported that nonlactating bison (Birn bison) apparently bredearlier than did cows thatwere

nurSing.

Comparison of median dates of conceptionwithin the respective November and post November collectionsof adult Roosevelt and Rocky Mountain elk (Tables 1 and 2)suggests that the rut commenced earlier in 1968 than in 1967. An obviousexplanation is that cows were

in better physical condition during thelate summer of 1968 than fora similar period in 1967.Unfortunately, measurements of physical condition (available only for Roosevelt elk)during the two reproductive years were recorded at different chronologicalseasons and are therefore not comparable. Nevertheless, otherevidence is available which indirectly suggests an improved nutritionalstatus for cows during the prerutting period of 1968.Weather conditions prevailing during the

summer of 1967 as compared to 1968 were strikinglydifferent,The June to mid-September interim of 1967was one of the driest on record for Oregon, and observations indicatedthat plant growth was severely retarded as a result,Although little precipitation fell from June to August 10, 1968, unprecedented rainfalloccurred subsequently in August throughout Western and EasternOregon, and the production of forage responded accordingly.It appears, therefore, that earlier breeding activity apparent for adult femaleRoosevelt and Rocky 20 Mountain elk in1968resulted from improved physical condition because of greater availability of forage during the prerutting period.

Fertility

Pregnancy Rates

Prior to commencement of the present study itwas speculated that the disparity of pregnancy rates observed between samples of adult Roosevelt and Rocky Mountain elk was largely attributable to collection of female Roosevelt elk too early in the season.Possibly some cows were examined before they were physiologically ready to ovulate, or more likely, before all embryos actually presentwere large enough to allow detection of pregnancy.Three types of evidence were used to estimate the extent that time of collection influenced pregnancy rates observed for collections of Roosevelt elk examined during the five reproductive seasons(1964-68)included in this study (Table 3).

(1) Of the95adults collected on all areas during1965, 1967, and1968and designated as nongravid, both ovaries of79were sectioned,Sixty-eight(86percent) either lacked corpora lutea, contained regressing corpora lutea, or were from uteri containing non-viable embryonic tissue.Therefore these cows could not have beenpreg- nant when killed.The remaining 11 cows with functional corpora lutea were distributed in the three samples as follows,1965- two, Table 3.Pregnancy rates of Roosevelt elk collected in Western Oregon, 1964-68 reproductive seasons (collection dates in parenthesis). No. Examined No. Pregnant Pèrcent Pregnant Area and Reproductive SeasonYearling Adult Yearling Adult Yearling Adult

Millicoma / 1964 (Nov. 28&29)- 0 46 - 25 - 54 1965 (Nov. 20&2l)/ 5 41 0 ' 10 0 24 1966 (Dec.3 & 4)1 7 38 1 22. 14 58 1967 (Jan. 3 Feb. 25, 68) 1 48 0 24 0 50 1968 (Nov. 2 Dec. 14) 4 50 0 31 0 62. TOTALS 1964-68 17 2.23 1 112 6 So

T T

1968 (Nov. 17Dec. 15) 3 21 1 10 33 48 North Coast 1967 (Nov. 11 - 18) 2. 12 0 5 0 42 1968 (Nov. 16, 68Feb,15,69) 3 8 1 5 33 62. TOTALS 1967-68 5 20 1 10 20 50

Totals All Areas 1964-68 2.5 2.64 3 132. 12. 50

Harper, 1965: 7 Harper, unpublished field notes, 1965 Harper, 1967:3 22

1967 - four, and 1968- five.Even if all 11 actually were gravidor were about to ovulate (a doubtful assumption since allwere collected after November 1), changes in the respectivereproductive rates would have been minor. Both ovaries from 14 of 16nongravid yearlings killed during 1965, 1967, and 1968were also examined. None con- tamed corpora lutea.

(2) Comparison of breeding dates for Rooseveltelk collected during the November and post November periodsof the 1967 and 1968 reproductive seasons combined (Table 1) suggests that ifall specimens obtained in November had been collected in Decemberor later, the pregnancy rates for samples collected in November would have been about six percent higher. (3)Data on dates of breeding or frequency of ovulationare lacking for the 1964 and 1966 samples of Roosevelt elk.But 54 and 58 percent of the adults included in these samples collectedon November 28 and 29, 1964 and December 3 and 4, 1966,respectively were gravid, compared to a 50 percent pregnancy rate for the sample ofadults examined during January and February, 1968 (Table 3),If it were true that many pregnancies went unnoticed in uteri collected inlate November and December of 1964 and 1966, respectively,than a lower fertility rate would hardly be expected fromcows autopsied beginning about three weeks later during January. The foregoing evidence suggests thatsome potential Z3 pregnancies" were not enumerated in Roosevelt elk examined in November, But the number involved was evidently small and hardly sufficient to account for the marked difference in reproductive rates apparent between Roosevelt and Rocky Mountain elk. One-half, or 13Z of Z64 adult Roosevelt elk checked from 1964 to 1968 were gravid (Table 3).The average rates of pregnancy for adults from each study area were almost identical, but greater variation appears in ratios of pregnancies observed annually for the respective localities. Among the ratios of gravid adults recorded for each reproductive season on the Millicoma area, only the proportion of pregnancies found in 1965 is indicated to differ significantly from the mean of 50 percent for the area (chi-square14.93, 4 df--most of difference, 10.95,attributed to 1965 season).As compared to other seasons, reproductive rates among adults in 1965 were substantially lower for all ages represented. Furthermore the calf: cow ratio recorded in March, 1967, which reflects the 1966 birthrate corresponding to the 1965 season of reproduction, was only 30: 100 contrasted to an average of

37: 100 for the reproductive years, 1964-67 (Harper, 1969: 2).Con- sequently, the estimate of pregnancies for 1965 evidently reflects a change in reproduction (for unknown reasons) rather than age-related differences in fecundity or random variation. A ZO percent difference in fertility levels for the 1967 and 1968 24 collections of adult females from the North Coast area (Table 3) is indicated, but the dissimilarity is possibly due to the small number of elk studied, The age specific pregnancy rates of Roosevelt elk studied are corn- pared in Table 4 with similar data recorded for certain other populations of elk,Of the Roosevelt elk examined, cows from four to ten years old had the highest pregnancy rates. Few yearlings were gravid, and none of eight cows judged to be older than tenyears were pregnant.The evidence of a difference in the proportions of pregnancies found among the two and three year olds as compared to the four to ten year olds was significant (chi-square = 12. 34, 1 df). The pregnancy rates shown for Rocky Mountain elk collected in Northeast Oregon are based on cows obtained during the November- January periods of the 1967 and 1968 reproductive seasons. Compari- son of breeding dates for adults killed during the November and post November periods of both seasons (Table 2) and comparison of pregnancy rates for 68 adults collected in November (85 percent) with the 95 percent rate for 22 adults killed in December and January mdi- cates that about 10 percent more pregnancies would have been apparent had all uteri been collected beginning in December, The proportions of pregnancies indicated for all age classes of Roosevelt elk except yearlings are lower than those found in all but one comparable age group of Rocky Mountain elk (Table 4),This Table 4.Comparison of age-specific pregnancy rates among samples of Roosevelt and Rocky Mountain elk (outside collection dates in parentheses-- all years listed as reproductive seasons). All Age in Years 1 2 3 4 5-7 8-10 11+ Ages 2+

Roosevelt Elk (This Study) Exam. 25 27 33 49 74 22 8 238' 213 1965_6&U Preg. 3 9 10 29 43 13 0 107 104 (Nov. 2-Feb. 25) °/ Preg. 12 33 30 59 58 59 0 45 49 Rocky Mountain Elk Northeast Oregon Exam. 15 13 23 22 14 16 2 105 90 (This Study) Preg: 5 12 19 19 14 13 2 84 79 1967&1968 (Nov. 2Jan. 25) %Preg. 33 92 83 86 100 81. 100 80 88

N. Yellowstone Exam. 104 79 56 62 141 78 74J 584 480 (Greer, 1966) Preg.1/ 1963-1966 12 75 53 61 138 77 59 475 463 (Oct. 27June) %Preg. 12 95 95 98 98 99 80 81 96 Michigan Exam. 11 21 27 17 20 6 3" 105 94 (Blouch&Moran, 1965) Preg. 2 16 20 14 1964 17 2 2 73 71 (Dec. 5-13) %Preg. 18 76 74 82 85 33 67 70 76

'Datafor 1965 and 1966 compiled from Harper,(unpubljshed fieldnotes, 1965) and 1967. V Less than the 289 females listed in Table 3 since 51 of the cows aged during the 1964-66 seasons were assigned to different age groupsthan I used. 'Computed fromnumber of specimens and percent pregnant. Classified as eight and nine year olds. Classifed as ten and older.

Ui 26 exception seems insignificant, since oniy six adults (ages eight and ten years, Michigan) are involved.Though a trend is apparent in collections of Rocky Mountain elk to indicate that thepregnancy rates of cows four to ten years old are higher than in the otherages, the differences are much less pronounced than for Roosevelt elk. The incidence of pregnancies determined from adults in the three samples of Rocky Mountain elk listed in Table 4 approximate the 74 to 94 percent fertility rates that Kittams (1953: 181) summarized for 11 samples totaling 2, 072 adult cows collected intermittently from 1935 to 1950 in Yellowstone National Park, Jackson Hole, and Banff National Park, Canada.Therefore the 50 percent pregnancy rate calculated for adult Roosevelt elk in Western Oregon (averagepropor- tion of gravid adults in Tables 3 and 4) is only one-half to two-thirds the rates reported for adult cows from other elk herds in North America.

Incidences of Twinning and Multiple Ovulation

Of 135 pregnancies in Roosevelt elk (Table 3), and 84 in Rocky Mountain elk (Table 4) checked in my study, all involved only single embryos. Kittams (1953: 183) reported only five twinningsamong

1,690 gravid Rocky Mountain elk uteri. Both ovaries of 83 gravid Roosevelt and 68 gravid Rocky

Mountain elk in the above samples were sectioned.Although the Z7 ovaries of no pregnant cow contained more than one corpus luteum over 11 mm in diameter, 78 percent of the gravid Roosevelt elk and 85 percent of the gravid Rocky Mountain elk had developed secondary corpora lutea (ranging from 3 mm to 10 mm in diameter), which appeared functional and apparently were the result of postconception ovulations (Halazon and Buechner, 1956 and Morrison, 1960).

Evidence of Infertility

I conducted an intensive laboratory study of 75 nonpregnant Roosevelt elk uteri collected during the 1967 and 1968 reproductive seasons, and Z8 nongravid uteri obtained in 1965 to determine in what manner infertility was manifested.

Infertility in Yearlings

Sectioning of both ovaries of 14 nonpregnant yearlings revealed no corpora lutea or corpora albicantia (pigmented scars that deve'1op from corpora lutea of previous cycles).If the sample of ovaries examined is typical, it appears that the low incidence of pregnancy apparent for yearling Roosevelt elk (1Z percentTable 4) is because most yearlings do not attain sexual maturity by the second rutting season of their life. 28 Infertility in Adults

The infertility identified in adultcows was evident as a failure to ovulate and as embryonic mortality(Table 5). Anovulation.Both ovaries of 79 nongravid adultswere examined. At least 18 had not ovulatedduring the reproductive season of their collection.Except for one cow with cystic ovarian follicles, the ovaries of femalesconsidered to be anovulatory

appeared normal but lackedcorpora lutea or corpora albicantia,Of the remaining 61 pairs ofovaries, 29 had corpora lutea and 32 showed only corpora albicantia,

My conclusion that 17cows with seemingly normal ovaries failed to achieve estrus is basedon the assumption that all corpora albicantia originated from luteal tissue of thesame breeding season as when the female was killed.Morrison (1960: 306), who studied ovaries from elk of known breeding history, indicatedthat many corpora albicantia were retained for more than one year, and that hewas unable to dis- tinguish pigmented scars of previousreproductive seasons from those of a current season,It appears, then, that some of the 32 adults whose ovaries showed pigmentedscars but no corpora lutea might not have ovulated within theyear.Therefore I consider the 18cows listed as anovulatory in Table 5 to bea conservative figure. A more realistic assessment of the proportionof nongravid adults in this Table 5.Infertility identified in nongravid adult Roosevelt elk collected in Western Oregon, 1965, 1967 and 1968reproductive seasons. Infertility Identified

Uteri Possible Embryo Ovulation' Ovulation" Anovulation Mortality Total

Collection Period Area Exam.1 No. No. No. °, No. No. November

20-21; 1965 Millic. 20 7 - 8 - 5 - 0 - 5 11-18, 1967 N. Coast 5 3 - 1 - 1 - 0 1 1 16-17, 1968 N. Coast 3 2 - 1 - 0 - 0 - 0 2-22, 1968 Millic. 18 7 5 5 1 - 6 - December-February

1/3/68-2/25/68 Millic. 21 2 12 7 0 - 7 12/7-12/15/68 LoonLk. 12 6 - 5 0 1 - 1

Totals by Age Groups 2-3 yrs. 23 8 35 7 30 8 35 0 0 8 35 4-lOyrs. 46 18 39 22 48 5 11 1 2 6 13 11 + yrs. 10 1 10 3 30 5 50 1 10 6 60

Adults 79 27k' 34 32 41 18 23 2 2 20 2.5

'Uteriwith both ovaries present. V.With corpora lutea. With corpora albicantia. Without corpora lutea or corpora albicantia. 'Includestwo cows listed as "aged" in 1965 collection "Specimenswith dead embryos omitted from total. 30 sample that did not ovulate, more likely should liesomewhere between 23 and 64 percent (the percentage withoutcorpora lutea or pigmented

scars plus the percentage of ovaries containing pigmentedscars only.-- Table 5).

Embryonic Mortality.Early embryonic death was readily apparent in two of the 79 reproductive tracts of nonpregnantadult Roosevelt elk (Table 5).I consider the incidence of intrauterine mortality given as minimal for tworeasons.First, of the 20 uteri collected in 1965 that I examined, most had beenpreviously opened in the field to check for possible pregnancies.Because of the possi- bility that any embryonic remains present mighthave been carried out with fluid released when the uteri were first incised, thechance of detecting prenatal losses in these specimenswas reduced. Second- l.y, early embryonic mortality proved difficult to determine because circumstances concerned with collection of reproductivetracts pre- cluded a thorough examination of uterine contents whilefresh.Most nongravid uteri obtained in 1967 and 1968were placed in dry ice or in- jected with and then placed in AFA withinone hour after autopsy to minimize post mortem changes.Though the latter method provided the best material for histological study, scrutiny of fresh,unfrozen specimens would have been more desirable.

The occurrences of prenatal mortality and suspectedmortality in all nonpregnant uteri examinedare described in Table 6.Chorionic Table 6.Description of occurrences and suspected occurrences of embryonic mortality for Roosevelt elk collectedin Western Oregon, 1965, 1967 and 1968 reproductive seasons. Collection Ovarian Examination* Age No. Date (Years) Udder Corpora Diameter Description of Uterine Contents

68-118 11-10-68 4 Wet lutea 13 mm (f) Mfsc2c: Fragmented remains of chorion (two largest pieces 4 mm (r) 8 mm x 55 mm); embryo not located.Histolog: Chorionic albicantia 3 mm (2) epithelium undergoing extensive degeneration.Embryo probably died at about 20-25 days.Diagnosis: Embryonic tissue dead.

68-132 12-7-68 11-15 Wet lutea 13 mm (f) Macroscopic: Two frayed pieces of chorion (8 mm x 15 mm and albicantia 2 mm (3) 4 mm x 11 mm); Histolog: Tissue organization lacking, dead 3 mm (2) chorionic cells.Diagnosis: Embryonic tissue dead.

67-141 11-18-67 5 (one ovary lost in coll.) Macroscopic: Compact body of tissue (12 mm x 6 mm x 3 mm) lutea in tip of uterine horn.Histolog: Tissues poorly defined, but aggregations of nucleated cells the same size as embryonic red albicantia 4 mm blood cells of healthy chorion apparent.Diagnosis: Embryonic tissue dead.

67-62 11-11-67 3 Dry lutea 10 mm (r) Macroscopic: Membranaceous fragments 1 mm to 2 mm in size. albicantia 4 mm Histolog: Many necrotic cells resembling chorionic cells. Diagnosis: Embryonic deathsuspected.

67-93 11-11-67 3 - lutea 12 mm (f) Macroscopic: Membranaceous fragments 1 mm to 3 mm in size. 4 mm (r) Histolog: Many necrotic cells resembling chorionic cells. albicantia 2 mm (2) Diagnosis: Embryonic death suspected.

* (f) and (r) = corpora lutea appearing as functional and regressed, respectively;number of corpora albicantia in excess of one enclosed by parenthesis.

tJ-

1 32 vesicles of about 20 to 25 days of age were found broken in uteri of two additional three year old cows collected, November, 1968,I could not find an emLryo in one of the chor ions, and the 6 mm embryo in the other uterus looked less developed than 6 mm embryos associated with normal appearing membranes. Though some fragmentation of tissues was evident in histological sections of both vesicles, the structural distortion was less than observed in tissues described in Table 6.Considering the absence of more definite evidence of tissue degeneration, and because intrauterine injection of preserving fluid can rupture embryonic membranes(B0W. OGara, personal communication, 1969), both specimens were considered pregnant. As was probably true for anovulation, the incidence of early prenatal deaths in Roosevelt elk studied is probably greater than the two percent figure listed in Table 5.

Infertility in Rocky Mountain Elk

Of 17 nongravid Rocky Mountain elk collected in Northeast Oregon during the 1967 and 1968 reproductive seasons, the ovaries of two of eight yearlings and eight of nine adults examined contained corpora lutea (Table 7).A large corpus albicans (4. 5 mm) was observed in the ovaries of the adult specimen that lacked luteal tissue, thus suggesting that this cow had also ovulated during the current breeding season,Nonviable embryonic tissue (similar circumstances Table 7. Infertility identified in nongravid Rocky Mountain ell collectedin Northeast Oregon, 1967 and 1968 reproductiveseasons.

Infertility Identified Possible Embryo Uteri Ovulation' Ovu1ation Anovulation1 Mortality Age Collection Period Exam)' No. No. No. No. Total No. Yearlings Nov.11-19, 1967 4 0 0 4 0 4 Nov.17, 1968 2 0 2 Totals 8 2 0 6 0 6 Adults

2-3 yrs. Nov. 1 12, 1967 0 1 0 0 0 Nov.16, 1967 Dec.22, 1968 3 2 0 0 1 1 4-10 yrs. Nov. ii, 1967 3 3 0 0 0 0 Nov. 16, 1968 2 2 0 0 0 0 11+yrs. _. Totals 9 7k" 1 0 1 1 UUteri with both ovaries present. ai. With corpora lutea. 'Withcorpora albicantia. 'Withoutcorpora lutea or corpora albicantia. "Appearanceof dead chorion similar to 68-132 (Table 6). Specimen with dead embryo omitted from total. 34 as for 68-132 -- Table 6) was found in one of the adults exhibiting a functional appearing corpus luteum, If the small sample of nongravid Rocky Mountain elk examined is representative, it appears that the frequency of anovulation, among adults at least, is much less than for the Roosevelt elk studied.

Factors Influencing Fertility

Pathology

Diseases Associated with Reproduction,Brucellosis, leptospirosis, and vibriosis cause substantial losses to reproduction in livestock, and acute infections of anaplasmosis in cattle frequently result in abortion (Gibbons, 1968:396). Rush (1932: 372) found evidence of brucellosis in elk from Yellowstone National Park, Serologi- cal reactors of Leptospira pomona have been found among white-tailed deer (Odocoileus virginianus) (Trainer and Hanson, 1960: 44), and anaplasmosis has been produced experimentally in white-tailed deer, black-tailed deer (Odocoileus hemionus columbianus), mule deer (0.Ii. hemionus), and Rocky Mountain elk (Howe and Hepworth,

1965: 114). Samples of blood obtained from 109 female Roosevelt elk killed or captured on the Millicoma area during January, February, March, and November, 1968 were tested for brucellosis (Brucella abortus), 35 leptospirosis (Leptospira pomona, L, canicola, and L. icterohaemorrhagiae), and Anaplasma marginale.All samples gave nega- tive reactions for brucellosis and leptospirosis.Additionally, vibriosis (Vibrio fetus var, veneralis) was not detected among 4Z cul- tures of cervical mucus collected from females killed in November, .: Almost80percent of the serum samples obtained from66 cows collected between January and March,1968reacted positively to a capillary tube agglutination test (CA) for the carrier state of anaplasmosis, Because of the high reaction rate and the failure to find the organism in red blood cells, Dr. Dean Smith of the Veterinary Diag- nostic Laboratory (Oregon State University) concluded that the serological test used was not valid for elk blood.Of 47 serum samples from cows killed in November,1968,only two, or four percent were positive for anaplasmosis. Since both collections of sera originated from the same localities and were collected and processed identically by the same technician, the results cast further doubt on the accuracy of the CA test for elk serum, Howe et al,(1964)has also reported on the inaccuracy of the capillary tube agglutination test as a method for diagnosing anaplasmosis in several species of wild ruminants, including elk, Abnormalities in Uteri and Ovaries The only abnormality recognized among uteri and ovaries of6 yearling andZZ9adult 36 Roosevelt and Rocky Mountain elk in which the uterus and both ovaries were available for study, was an occurrence of cystic ovarian follicles in an eight year old, nongravid Roosevelt elk that was dry when killed on the Millicoma area, Two cystic follicles measuring 40 mm x 40 mm and 4 mm x 20 mm occupied all of the right ovary, whereas the left ovary looked normal, and contained 13 follicles from 2mm to 8 mm in diameter and a 1.5 mm pigmented scar. The condition of the ovaries strongly suggested that ovulation had not occurred earlier in the season. When considering that cystic ovarian folliclesare corn- monly associated with impaired fertility in dairy cattle (Merck, 1967: 852), it seems likely that conception was prevented in this elk for the same reason.

Male Coverage

To measure the sufficiency of male coverage as related to fertility of females, calf: cow and bull: cow ratios during 1950-69were compared. Although essentially no change was apparent in thepro- portion of calves recorded throughout the 20 year period, the ratio of bulls observed from 1950 to 1959 is over twice the ratio from 1960 to 1969 (Table 8).The decline in bulls is related to substantial increases in numbers of elk hunters and regulation changes allowing harvest of yearling males.If the fertility of cows was depressed because of in- sufficient male service, then the apparent production of calves should 37 also drop with the decreased percentage of bulls.Because this has evidently not occurred, it appears that the present supply of bulls is ample for reproduction, and that the relatively low pregnancy rates observed in Roosevelt elk cannot be attributed to inadequate male coverage.

Table 8.Ratios of bulls and calves per 100 cows recorded annually for Roosevelt elk in Western Oregon between December and mid-February, l95O-7OJ/ Number Classified No. per 100 cows Period Cows Bulls Calves Bulls Calves

1950-59 7,964 1,704 3,215 21 40 1960-69 19,974 1,517 8,307 8 42

1950-69 27,938 3,221 11,522 12 41 l966-70' 14,470 701 5,962 5 41 Compiled from 1950-70 Annual Game Reports, Oregon State Game Commission, 7/ Period corresponding to reproduction seasons of study.

Physical Condition

Indices of Physical Condition,The physical condition of female Roosevelt elk collected during the 1967 and 1968 reproductiveseasons was measured to relate condition of the animal with fertility.Because of the limited time available for examination of elk in the field, and for other reasons to be discussed, indices of kidney fat,marrow fat (metatarsal) and adrenal weight were selected to assess physical i:i condition of the Roosevelt elk studied.(Measurements of fat depots and adrenal weights were not obtained from samples of Rocky Moun- tamelk,) Riney (1955) compared several indices for quantifying fat depots in red deer (Cervus elaphus) and documented the use of fat reserves as an indicator of condition for several species of animals, He states (p. 430-431) The fact that fat in ruminants appears to be largely endo- genous makes it a particularly useful index of metabolic level and potential energy reserves of the animal, and it is here postulated that fat can be taken as a direct measure of the condition, reflecting the metabolic level or goodness of physiologic adjustment of an animal with its environment. Of the indices he studied, Riney concluded that the kidney fat index (weight of the fat around the kidney expressed as a percentage of the weight of the kidney) best reflected seasonal changes in fat reserves among both sexes of red deer.Riney (1955: 435) found that fat depots in femur marrow apparently were not mobilized until after depots of subcutaneous and visceral lipids were mostly exhausted. Consequently, variations in the fat content of femur marrow occurred only for deer in the lower half of Riney's scale of ideal condition (p.

446). Ransom (1965) compared the percentage of fat in femur marrow with kidney fat indices of 34 female white-tailed deer killed between December and April in Manitoba, Canada.His data showed that the 39 kidney fat index decreased to about 30with little change in fat content of femur marrow, but after this pointwas reached, femur fat decreased markedly. Ransom also found thatdecreases in kidney fat when indices fall below 30were irregular relative to percentages of femur fat,He concluded that the kidney fat indexwas a satisfactory indicator of physical condition at valuesat or above 30, but recom- mended using femur fat toassess condition when kidney fat is less than 30. Anderson, Medin, and Ochs (1969:337) estimated the percent of fat in carcasses of 18 wintering mule deerusing seven different indices, and concluded that the kidney fatindex was probably the most useful if based on sufficient sample size.They reported a correlation coefficient of 0.67 between percent kidney fatand percent ether- extractable fat in the carcasses of deer examined. The relative size of the adrenal glands isconsidered an indicator of stress in various species of mammals includingdeer (Hughes and Mall, 1957; Welch, 196Z).Christian and Davis (1955: 177) believe that adrenal cortical hormonesare mainly involved in counter- acting the effects of stress stimuli, and thatchanges in the amount of adrenal cortex are related to the quantity ofcortical hormones produced.Therefore the amount of adrenocortical tissuelike the size of fat reserves should provide measurement ofthe physical condition of the animal.

Yearlings, two years old, andcows older than ten years are 40 considered separately from other females in comparisonsof physical condition and fertility.Because all pregnancies in yearl&ngs and many in two year olds were associated with puberty (Table 4), lactation and other physiological conditions resulting fromprevious pregnancies would be absent as factors affecting conceptionin these ages. Similarly, cows older than ten years (all nonpregnant)are excluded from comparisons of animal condition and reproductionsince factors concerned with senility might be responsible for theirnongravid status.

Indices of Kidney and Marrow Fat.The kidney fat indices and values of percent ether-extractable fat in metatarsalmarrow of 95 female Roosevelt elk, three years and older, collectedduring the 1967 and 1968 reproductive seasons are shown in Figures4 and 5.Meta- tarsal marrow was analyzed instead of femurmarrow because it was impractical to remove samples of femurmarrow from elk shot by hunters,Both plots indicate the same general relationshipbetween kidney fat and the fat content of metatarsalmarrow as Ransom (1965) described for kidney and femur fat in deer, When the reproductive condition ofcows (3-10 years old) is considered relative to fat stores it is apparent thatpregnant females had larger deposits of kidney andmarrow fat than nonpregnant cows

(Figures 4 and 5).Females with a kidney fat index above 60 hada 93 percent pregnancy rate, those with an index between 59 and 30showed I., A S ® .®: .. 80

4OL©

I = Pregnant (Nonlactating) A Nonpregnant (Nonkctating) 20 o= Lactating -= 11+years

40 80 120 160 200 240 280 Kidney Fat Index Figure 4. Kidney fat indices plotted against percentage of fat(wet basis) in metatarsal marrow for 54 femaleRoosevelt elk, 3 years and older, collected in WesternOregon, November 2 - December 15,1968. 1968. 28, 3 Oregon, in 3 female120 41 for marrow 100 metatarsal- January in basis)80 (wet Western Index fat of Fat peentage60collected Kidney againstolder, and 40plotted years indices elk, fatRoosevelt 20 Kidney 5. Figure (NOfliactating) Nonpregnt yearsLactating + LI -0A 20 (Nonlactating) Pregnant 60 S 80 . . A A A 1 43 a 50 percent pregnancy rate, and only ten percent of the females with kidney fat values under Z9 were gravid. Low reproductive rates also coincided with percentages of fat in the metatarsal marrow under about 85. Using stepwise multiple linear regression, kidney fat indices of 89 females, 3-10 years old, listed in Figures 4 and 5 were compared against effects of age, reproductive status, status of lactation and month of collection.Because elk were collected on the three study locations at different periods during each reproductive year, compari-

Sons of fat reserves between areas or years were not possible.The influence of age on kidney fat deposits was negligible (p > 0. 50), but gravid cows has significantly greater fat depots than nongravid cows (p < 0.001), and lactating females showed significantly less kidney fat than nonlactating cows (p < 0.001). Measurements of kidney fat deposits were obtained from eight yearlings and nine two year old Roosevelt elk.Though the sample is small, gravid females had higher average indices of kidney fat than nongravid females (Table 9). Lactating cows had lower pregnancy rates than nonlactating females in the collections of Roosevelt elk examined.In a sample of 60 cows, 3-10 years old, collected during November and December, 1968 for which data on lactation are available, 51 percent of the wet cows were gravid compared with 84 percent of the dry females Table 9.Indices of kidney fat for yearling and two year old female Roosevelt elk collected in Western Oregon, 1968 reproductiveseason; all specimens nonlactating.

Kidney Fat Indices: (Number Cows in Parenthesis) Yearlings Two Year Olds Pregnant Nonpregnant Pregnant Nonpregnant Collection Period Range Mean Range Mean Range Mean Range Mean

November 2-17, 1968 (0) - - (6) 35-123 71 (3) 101-198 152 (5) 73140 110

December 7-15, 1968 (1) - 96 (1) - 19 (1) - 175 (0) - 45 (Table 10). A chi-square test showed that the evidence ofa difference in pregnancy rates between wet and dry cows was significant (chi-square = 6.80, 1 df). The proportions of lactating cows in samples of Roosevelt elk (3-10 years old) collected in November, December andJanuary, 1968 were: 24 of 42 (57 percent), 11 of 18 (61 percent), and 17 of 42 (40 percent), respectively (Table 10).The similar percentages of wet cows recorded during November and December, 1968 as compared to the substantially lower incidence of lactation observed in January, 1968 suggests that many females ceased lactating inJanuary. Because gravid and nongravid females might not 'dry up" at similarrates, comparison of pregnancy rates between wet and drycows are probably least biased for specimens obtained during November and December. Examination of udders of 65 Rocky Mountain elk, three to ten years old, collected during November and December, 1967 and 1968 also revealed a lower incidence of pregnancyamong wet cows (82 percent) as contrasted with dry cows (100 percent-- Table 11).But the indicated difference in pregnancy rates between lactating andnon- lactating cows (18 percent) was not significant (chi-square= 2.00, 1 df) and is considerably less than the 33 percent difference recorded for Roosevelt elk collected in November and December, 1968 (Table 10). The comparison of fat deposits with reproductive status in Table 10.Relationship of pregnancy rates and status of lactation forRoosevelt elk, 3-10 years old, collected in Western Oregon, 1967 and 1968 reproductiveseasons. Lactating Nonlactatirig Number Number Number Number Collection Period Examined Pregnant Examined Pregnant 1967 Reproductive Season

January 3-28, 1968 17 7 2.5 16 1968 Reproductive Season November 2-22, 1968 2.4 13 18 16 December 7-15, 1968 11 5 7 5 Totals by Months November-December 35 18 25 21 Percent Pregnant 51

January 17 7 25 16 Percent Pregnant 41 64 Totals 1967 and 1968 Reproductive Seasons 52. 25 50 37 Percent Pregnant 74

0" Table 11.Relationship of pregnancy rates and status of lactation for Rocky Mountain elk, 3-10years old, collected in Northeast Oregon, 1967 and 1968 reproductiveseasons.

Lactating NonLac tat ing Number Number Number Number Collection Period Examined Pregnant Examined Pregnant 1967 Reproductive Season November 11-19, 1967 21 16 4 4 December 11-22, 1967 3 3 1968 Reproductive Season November 2-20, 1968 26 22

December 7-31, 1968 6 5 3 3 Totals 1967 and 1968 Reproductive Seasons November-December 56 46 9 9 Percent Pregnant 82 100

A Roosevelt elk studied reveals a positive relationship betweenphysical condition and fertility.The relationship was apparent in females with pregnancies less than 30 days advancedas well as in cows during later stages of gestation.Therefore it is doubtful that the larger fat stores typical of gravid cows accumulatedas a result of physiological factors associated with pregnancy. More likely, thedifference observed in fat depots between pregnant andnonpregnant Roosevelt elk (3-10 years old) reflects factors suchas lactation which affect the female before conception.

Indices of Adrenal Weight.Hughes and Mall (1957: 194) found that adrenal weights of Z7 adult female black-taileddeer (Odocoileus hemionus columbianus) collected during early Novemberin California were reasonably well correlated (r = -0. 57) with four classes of physical condition (based on determinations of kidney fat).The animals they studied were from a population believedto be in excess of the carrying capacity of the habitat.They concluded (p. 195) "that in the deer sampled adrenal cortical size, most easily measuredas adrenal weight, may be considered asa condition factor with regard to the field condition determinants presently inuse, " The relationships between adrenal weights in 76 female

Roosevelt elk, 3-10 years old, collected during the 1967and 1968 reproductive seasons and variables ofseason, age, pregnancy, lactation and percent metatarsal fatwere analyzed using stepwise multiple 49 linear regression. Simple coefficients of correlation resulting from the analyses are presented in Table 12.The direction (sign) of the relationships of adrenal weight versus variables of fat depots, pregnancy and lactation agree with interactions previously described between fat reserves, pregnancy and lactation.The strong positive correlation between unadjusted adrenal weight and age probably reflects the effect of body weight on adrenal size, since the weight of cows in the 3-10year age group also generally increased with age (C. E. Trainer, unpublished data, 1969). Of the three indices used to express adrenal size (Table 12), relative adrenal weight expressed as centigrams of adrenal tissueper pound of hog-dressed carcass weight generally resulted in the strong- est correlation with other variables, except for age.(Unless otherwise stated, relative adrenal weight as used in reference to Roosevelt elk will refer to the adrenal weight: hog-dressed weight ratio. The quantitative importance of the independent variables shown in Table 12 to relative adrenal weight in the stepwise regression is indicated by the R2 values in Table 13. Among the variables analyzed during both seasons, only lactation, which accounted for 32 percent

(R2x 100) of the variation in relative adrenal weight in the January, 1968 sample, substantially influenced adrenal size.The relationships shown were not improved by similar analysis using kidney fat indices Table 12.Correlation coefficients (r) computed from correlation of three indices of adrenal weight withvariables listed for female Roosevelt elk, 3-10 years old, collected on Millicomaarea, 1967 and 1968 reproductive seasons.

Percent Fat Percent Fat Reproductive Kidney Metatarsal Metatarsal Adrenal Weight Fat Season' lnd1ces/ Marrow Marrow Me Pregnancy Lactation Index (Wet Basis] (Dry Basis) g Adrenal 1967 A (Unadjusted wt) 0. 65 -0.08 0. 38 -0.20 -0.27 -0. 22 (Jan. 3-28, 1968) cg Adrenal per B lb Body wt 0.47 No. Examined 37 -0.01 0.45 -0.27 -0.36 -0.23 eg Adrenal per lb Hog-dressed C Body wt/ 0.53 -0.04 0.56 -0.32 -0.40 -0. 29 1968 A 0. 42 -0. 13 0.02 -0.07 -0.02 -0. 19 (Nov. 2-Dec. 14, 1968y" B 0.22 -0.27 0.06 -0.21 -0.12 -0.03 No. Examined 39 C 0.24 -0.27 0.13 -0.28 -0.14 -0.01 1967-1968 A 0.53 -0.05 0. 17 -0. 16 -0.22 -0. 18 No Examined 76 B 0.34 -0. 16 0. 22 -0.27 -0.32 -0. 23 C 0.37 -0.18 0. 30 -0.36 -0.39 -0. 30 'Collectiondates in parenthesis. 'Basedon total weight of both adrenals.

'Hog-dressedweight is carcass weight with all viscera removed,but with head, hide and feet attached. 'Thirty-sevenspecimens collected November 2-16; remainingtwo collected December 14. 51

Table 13.Coefficients of determination (R2) and F values resulting from stepwise multiple linearre- gression analysis of adrenal weight:hog-dressed body weight ratio Y on variables listed for female Roosevelt elk, 3-10 years old, collected on Millicomaarea, 1967 and 1968 reproductive seasons. Reproductive Array of Cumulative Individual 2/ Season VariablesX R2 -'1' R2 1967 Lactation 0.315 0.315 18.49 (Jan. 3-28, 1968) Age 0.462 0.147 8.65 No. Examined 37 Pregnancy 0.482 0.019 1 13 KidneyFat 0.487 0.005 0.31 Percent Fat Metatarsal Marrow (Wet) 0. 487 0.003 0.02 Percent Fat Metatarsal Marrow (Dry) 0.489 0.001 0.01 1968 Kidney Fat 0.081 0.081 3.18 (Nov. 2-Dec. 14) Age 0.123 0.042 1.67 No. Examined 39 Pregnancy 0.158 0.034 1.35 Percent Fat Metatarsal Marrow (Wet) 0.170 0.013 0.49 Percent Fat Metatarsal Marrow (Dry) 0. 178 0.008 0.31 Lactation 0.186 0.008 0.31 1967-1968 Percent Fat No. Examined 76 Metatarsal Marrow (Wet) 0. 155 0. 155 15. 85 Age 0.264 0.109 11.23 Year 0.301 0.037 3.83 Lactation 0.332 0.031 3.14 Percent Fat Metatarsal Marrow (Dry) 0.336 0.004 0.45 Kidney Fat 0.337 0.000 0.04 Pregnancy 0.337 0.000 0.00

x 100 = percent of variation in dependent variable Y that is attributable to combined effects of independent variables X. = ratio of individual R2/df to residual of individual R2/df. 'Thirty-sevenspecimens collected November 2-16; remaining two on December 14. 52 truncated down to 40 to remove the affect ofvery fat cows upon relative adrenal weight. Consequently, it appears that adrenal size,as measured by weight at least, in Roosevelt elk examinedwas largely determined by factors not included in the regression analyses. 53

IV.DISCUSSION

The pregnancy rate of 50 percent determined foradult Roosevelt elk examined apparently represents the fertility levelin adults during approximately the initial three months of gestation (throughJanuary). Knowledge concerning the extent of prenatal lossesbetween January and time of parturition would be desirable, especiallysince there are conflicting reports about the incidence of fetalmortality in elk during the late winter and spring.

Greer (1966: 124) mentions that the difference betweenan 89 percent rate of pregnancy for 642 adult Rocky Mountain elk collectedfrom the northern Yellowstone herd, November 21, 1961-February15, 1962 and an 80 percentpregnancy rate for 89 adults killed February, 21 to June 12, 1962 might suggest that intrauterine lossesoccurred. But he further states (ibid)

that evidence of resorption or recent abortionwas not found in any of the spring specimens. The lower fertilityrate was due to the large representation of older, and usually barren, females in the sample. Nearly 50 percentwere eight years or older. Murie (1951: 142), however, reports that althoughpregnancy rates of about 90 percent prevailed among large samples of adult RockyMoun- tam elk killed in the Jackson Hole-Yellowstone National Parkarea during December and January, 1935-36 and January, 1943,examination of hundreds of elk that died during the late winter indicatedthat 54 about 50 percent of the adult cows were nonpregnant. He concluded (ibid) that It is true that these were animals that had died, and that the same proportions may not have applied to the rest of the herd, yet this pregnancy rate may have been significant.If so, then it is a fact that about hail thecows do not produce calves in the spring.This is affirmed to some extent by field observations in the spring. Although only three female Roosevelt elkwere collected after

January (one 10-15 year old and one yearling, February 25,1968 -- both nongravid, and one gravid yearling, February 15,1969),prenatal losses occurring in Roosevelt and Rocky Mountain elk after about January were calculated using an indirect method.

The presence of milk in an udder is evidence that thecow was nursing during the period before she was collected.Because substantial numbers of calves are apparently not weaned until afterDecember, the incidence of lactation during November and December shouldserve as a measurement of the proportion of cows with calves at heel during that time of the year.For Roosevelt elk, the pregnancy rate of 55 percent for 44 cows, 2-10 years old, examined on the Millicomaarea, January,1968closely approximates the 57 percent incidence of lactation recorded for 42 cows, 3-10 years old, collectedon the same area the following November and December. Similarly, thepregnancy rate of87percent for 39 Rocky Mountain elk, 2-10 years old, killed in

Northeast Oregon, November,1967to January,1968was identical to 55 the 87 percent lactation rate recorded for 32 cows, 3-10years old, obtained during November and December, 1968 throughout thesame region. Though some mortalities of fetuses and/or calves undoubtedly

occurred in Roosevelt elk between January and November, 1968,none were indicated in the comparison of indices of pregnancy and subsequent lactation.But, as 34 of 39 Rocky Mountain elk uteri examined were collected during November, and because it was indicated previously that about ten percent of the "potential pregnancies" in adult females of this subspecies killed in November were not enumerated, it is likely that a fertility rate greater than 87 percent prevailed in Rocky Mountain elk uteri examined in November. Thus some losses of fetuses and/or calves were indicated for this subspecies.The relationship apparent between pregnancy and subsequent lactation rates for both Roosevelt and Rocky Mountain elk suggests that incidences of pre and postnatal losses happening between about January and Novem- ber, 1968 among elk 2-10 years old were minor.Consequently, it appears that pregnancy rates determined for Roosevelt and Rocky Mountain elk during the December-January period, 1967-68 closely approximated the birthrate for 1968. The indicated decline in the proportion of bulls in Roosevelt elk herds (a result of increased harvests) coupled with no apparent decrease in calf: cow ratios (Table 8) strongly suggests that the relatively 56 low reproductive rate recorded for thesubspecies in western Oregon originated with the female.

None of 14 nonpregnant yearlings whoseovaries were examined had ovulated.The evidence of infertility observedin the sample of 79 nongravid adults intensively studiedwas apparent as embryonic mortality (minimum of two percent) and ovulationfailure (23 to 64 percent -- Table 5),The reason(s) for the nongravidstatus of about one- third of the cows in this sample whoseovaries contained corpora lutea are more difficult to explain.Since functional appearing luteal tissue was observed in 11 of the 27 nongravid specimens which hadovulated (Table 5), it is possible thata few of these cows actually were gravid, but that the embryo was insufficiently developedfor macroscopic detection.But it is also possible thatsome corpora lutea resulted from a silent heat period, in which ovulation occurred without normalbe- havioral estrus and subsequent copulation.Silent heat is common among farm mammals (Hafez, 1968: 324), and Simkin (1965: 744)presented data indicating that, apparentlybecause of silent heat periods, many ovulations occurring in moose (Alces alces) early in thebreed- ing season do not result in fertilization. Although data obtained inmy study do not rule out a greater occurrence of mortality of embryonic tissue (including phasescon- cerned with fertilization of ova) than recorded,the evidence cited above suggests that most of the nongravidRoosevelt elk examined did 57 not experience estrus, or at least what is consideredas a normal estrus. Considering that indices of physical condition reflect stresses

affecting the animal, the strong relationship apparent betweenphysical condition and fertility indicates that the lowpregnancy rates prevailing in Roosevelt elk examined resulted from the effect ofenvironmental factors adverse to fertility.Unknown factors might have been opera- ting to the detriment of physical condition.But the positive correlation between fat stores, which serve as energyreserves for the animal (Riney, 1955: 430), and fecundity suggests that theultimate cause of low fertility levels observed in Roosevelt elkwas the stressing effect of a nutritional plane inadequate for sustainingreproduction at a high level.In the discussion to follow it is understood that the food resources available to the animal, as well as utilization of nutritive stores within its body are subject to considerable modifications by weather conditions.

The effect of nutrition on reproduction in elk examinedwas manifested in different ways. As discussed, initiation of rutting activity was possibly influenced by forage conditions during 1967 and 1968.

The apparent manner in which the plane of nourishmentaffected fertility in female Roosevelt elk appeared tovary according to age groups.

Since corpora lutea or corpora albicantiawere not found in the ovaries of six of 11 two year olds or in two of ten three year olds examined, it is likely that a delay in puberty was responsible for the nongravid condition of about 38 percent (eight of Zi) of the two and three year old females studied. For sexually mature cows, the positive relationship revealed between fertility and physical condition, plus the significantly lower pregnancy rates indicated for wet cows strongly suggests that nursing resulted in a reduction of nutritive stores to a level apparently mimi- cal to reproduction.The effect of lactation on fecundity indicates a general pattern of every-other-year breeding in many adult female Roosevelt elk.That is, a cow which is suckling a calf during the summer will enter the breeding season with lower energy reserves than if she were dry.The chances of her conceiving are therefore lessened. But because of nonlactation during the succeedingsummer, her energy stores will be high in the fall and conceptionmore likely. The variation in percentages of gravid adults recorded annuallyon the Millicoma area, 1964-68 (Table 3) reinforces the observation that many cows of breeding age are pregnant only on alternate years.It appears that the pregnancy rates apparent in 1964 (54 percent), 1965 (Z4 percent) and 1966 (58 percent) reproductive seasonson the area reflected such a breeding pattern, presumably as a result of factors inimical to the physical condition of the female (hence reproduction) during 1965.(A chi-square test comparing pregnancy rates for the 59 two reproductive seasons of lowest fertility (1965and 1967) with the three highest (1964, 1966 and 1968) indicatedthat the difference was significant - - chi-square= 8. 92, 1 df.) The proportion of adults breeding every-other-year is expectedto vary relative to the impact of such factors as weatherupon food supplies and stored energy reserves.

Since all of eightfemale Roosevelt elk over tenyears of age were nongravid (Table 4), it appears that reproductionin very old cows was particularly susceptible to the effects of impairednutritional intake.

The findings of this studyagree with results of other investiga-

tions concerning the influence of nutritionon fertility in cervids.The works of Buechner and Swanson (1955- - Rocky Mountain elk), Pimlott (1959-- moose), Daniel (1963 -- red deer) and Verme (1967-- white-tailed deer) all relate the importanceof food intake to attainment of puberty. Verme (1967) presented evidencedemonstrating that stress from lactation lowered fertility in white-taileddeer, and Lowe (1969: 440) found that thepregnancy rate of nonlactating red deer was mdi- cated as significantly higher than for lactatinghinds. Studies concerning reproduction in mule deer(Longhurst, Leopold and Dasmann, 1952: 57; and Julander,Robinette and Jones, 1961) and white-tailed deer (Verme, 1967)indicate that the nutritional level prior to and during the breedingseason was directly related to the rate of ovulation.The apparent effect of lactation upon fertility in Roosevelt elk clearly suggests that the months of June through November (period before and during the rut) are, from the nutritional standpoint, of critical importance to reproduction in this subspecies. However, food conditions during the winter are probably also significant to fecundity, especially as related to the onset of puberty, as has been indicated for moose (Pimlott, 1959: 398) and Rocky Mountain elk (Greer, 1966: 1Z8). When fertility rates of adult Rocky Mountain elk (88 percent pregnant) and adult Roosevelt elk (49 percent pregnant -- Table 4) collected in Oregon are compared in light of the indicated effect of nutrition on reproduction for the latter subspecies, it appears that Rocky Mountain elk were afforded a better nutritional regime than were Roosevelt elk.I believe this might be the case for two reasons. First, weather conditions prevailing in the habitat of the two subspecies are strikingly different,The climate of areas inhabited by Roosevelt elk in Western Oregon is characterized by mild tempera- tures, prolonged cloudy periods and heavy rainfall, which contrasts sharply to the warmer summers, colder winters and lower precipitation typical of the climatic environment affecting Rocky Mountain elk in Northeast Oregon.Therefore it is probable that weather conditions common to Western Oregon, particularly the higher amounts of rainfall and fewer days of sunshine, affect either the nutritive content of 61 forage eaten by Roosevelt elk, or in otherways stress the physiology of the female to the ultimate impairment of fertility.As winter

temperatires are milder and snowfall less in Roosevelt elk habitatof Western Oregon than on the Rocky Mountain elkranges of Northeast Oregon, one might conclude that weather conditions in thewinter should enhance physical condition in Roosevelt elk.Actually the con- trary might be true.Several stockmen with whom I have talked mdi- cated that it is more difficult to fatten cattle in thewet climate typical of the interim, October through March in Western Oregon,than in the colder but drier weather prevailing in Eastern Oregon duringthe same period.

No data are available on comparative population densitiesof Roosevelt and Rocky Mountain elk in Oregon.This is unfortunate since the higher fecundity typical of Rocky Mountain elk mightalso be the resuLt of greater abundance of forageper individual because of fewer elk per unit of habitat than for Roosevelt elk.Buechner and Swanson (1955) related an apparent increase in natalityamong yearling Rocky Mountain elk to greater availability of nutritionper animal because of a lower population density resulting from increased harvests. 62

V. CONCLUSIONS

Comparison of the average pregnancy rate recorded for adult Roosevelt elk examined during 1964- 68 reproductive seasons (50 percent -- Table 3) with the mean calf:cow ratio observed during the respective winters following these seasons (41 percent- - Table 8), indicates that the proportion of calves recorded was largely determined by the level of fertility rather than postnatal mortality.It is evident that the low fertility rates found in female Roosevelt elkwere not caused by inadequate male service or diseases.The apparent relationship between physical condition and pregnancy for cowsex- amined indicates that the fertility levels observed in female Roosevelt elk ultimately resulted from a lack of nutrition, or failure of the female to maintain nutritive stores necessary for a high rate ofrepro- duction.Lactation was identified to be the major factor affecting energy reserves of the cow to the detriment of fertility. Assuming that genetic differences among subspeciesare not responsible for variation in reproduction between Roosevelt and Rocky Mountain elk,it appears that the fertility rate observed in Roosevelt elk is limited either by (1) qualitative deficiencies in nutritionre- suiting from climatic influences or (2) by quantitative shortages in forage because of too great a density of elk.Therefore it seems important that measurements be obtained concerning the effect of various 63 population densities of Roosevelt elk upon fertility and net productivity the percentage of animals remaining after mortality from causes other than harvest has been deducted (Pimlott, 1959:393)].If fewer elk per unit of habitat would ultimately increase the number of animals available for cropping, then harvesting of antlerless elk would be justified.But if gains in net productivity did not result from lower population densities then the fertility level apparent for Roosevelt elk probably represents the maximum productivity possible under existing environmental conditions. 64 LITERATURE CITED

Anderson, A. E.,D, K. Medin and D. P. Ochs.1969.Relationships of carcass fat indices in 18 wintering mule deer.Proc. Western Assoc. State Game and Fish Commissioners 49:329- 340, Armstrong, R. A.1950.Fetal development of the northern white- tailed deer (Odocoileus virginianus borealis Miller). Am. Midi. Nat. 43(3): 650-666,

Batchelor, R. F.1965.The Roosevelt elk in Alaska, its ecology and management. Federal Aid in Wildlife Restoration Project W-6-R-5, Work Plan D.Alaska Dept. of Fish and Game. 3'7p. Blouch, R. I. and R. J. Moran.1965.1964 elk hunt.Res. and Development Rep. No. 30.Michigan Dept. of Conserv.24p. Buechner, H. K. and C. V. Swanson,1955.Increased natality resulting from lowered population density among elk in South- eastern Washington.Trans. N. Am. Wildi. Conf. 20: 560-567. Christian,J. J. and D. E. Davis.1955.Reduction of adrenal weights in rodents by reducing population size.Trans. N. Am. Wildl. Conf. 20:177-189. Daniel, M. J.1963.Early fertility of red deer hinds in New ZeaLand. Nature 200: 380.

Fuller, W. A.1966.The biology and management of the bison of Wood Buffalo National Park.Wildl. Mgmt. Bull. Ser. 1, No, 16.Canadian Wildl. Sev. SZp. Gibbons, W. J.1968.Viral, rickettsial and protozoan infections. In:Reproduction in farm animals (Ed. E. S. K. Hafez). Zd ed.pp. 381-401.Philadelphia, Lea and Febiger Co.44Op. Greer, K. R.1966.Fertility rate of Northern Yellowstone elk populations.Proc. Western Assoc. State Game and Fish Com- missioners 46: 123-128. Guyer, M. F.1953. Animal micrology.5th ed.Chicago, Univer- sity of Chicago Press,327p. 65

Hafez, E. S. E.1968.Reproductive failure in females.In:Re- production in farm animals (Ed. E. S. E. Hafez)pp. 321-341. 2d ed.Philadelphia, Lea and Febiger Co.44Op. haLCtzon, G. C. and H. K. Buechner,1956.Postconception ovulation in elk.Trans. N. Am. Wildl, Conf. 21: 545-554, Harper, J. A.1965.Ecological study of Roosevelt elk.Federal Aid ProjectW-59.-R-2,Job No. 1.Oregon State Game Com- mission.lOp. Harper, J. A.1967,Ecological study of Roosevelt elk.Federal Aid Project W-59-R-4, Job. No. 2.Oregon State Game Com- mission.3p.

Harper, J. A.1969.Ecological study of Roosevelt elk.Federal Aid Project W-59-R-6, Job No. 3.Oregon State Game Com- mission.7p. Howe, D. L., W. G. Hepworth, F. M. Blunt andG. M. Thomas. 1964. Anaplasmosis in biggame animals:experimental infec- tion and evaluation of serologic tests, Am, J,Vet. Res. 25: 1271-1275.

Howe, D. L. and W. G. Hepworth.1965.Anaplasmosis in big game animals:tests on wild populations in Wyoming. Am. J. Vet. Res. 26: 1114-1120.

Hughes, E. and B. Mall.1957.Relation of adrenal cortex to condition of deer.Calif. Fish and Game 44(2): 191-196. Julander, 0., W. L. Robinette and D. A. Jones.1961,Relation of summer range condition to mule deer herd productivity.J. Wildi. Mgmt. 2 5(1): 54-60. Kittams, W. H.1953.Reproduction of Yellowstone elk.J. Wildl. Mgmt. 17(2): 177-184,

Longhurst, W. M., A. S. Leopold and R. F. Dasmann.1952. A survey of California deer herds, their ranges and management problems.Calif. Dept. Game and Fish, Game Bull, No. 6, l36p.

Lowe, V. P. W.1969.Population dynamics of the red deer (Cervus elaphus) on Rhum.J. Anim. Ecol. 38(2): 425-457, Mace, R. U.1956.Oregon's elk.Wildi. Bull. No. 4.Oregon State Game Commission.33p. McCullough, D. R.1969.The tule elk, its history, behavior, and ecology.Univ. Calif. Pub. Zool. 88:1-191. Merck and Company.1967.The Merck veterinary manual.3d ed. Rathway, N. Jer., Merck and Co.l674p.

Morrison, J. A.,C. E. Trainer and P. L. Wright.1959,Breeding season in elk as determined from known-age embryos.J. Wildl. Mgmt. 23(1): 27-34. Morrison, J. A.1960. Ovarian characteristics in elk of known breeding history.J. Wildl. Mgmt. 24(3): 297-307. Murie, 0. J.1951.The elk of North America. Harrisburg, Pa. Stackpole Co. and Wildl. Mgmt. Inst., Washington, D. C.

Pimlott, D. H.1959.Reproduction and productivity of Newfoundland moose.J. WildI. Mgmt. 23(4): 381 -401, Quimby, D. C. and J. E. Gaab.1957.Mandibular dentition as an age indicator in Rocky Mountain elk.J. Wildl, Mgmt. 2 1(4): 435 -451.

Ransom, A. B.1965.Kidney and marrow fat as indicators of white- tailed deer condition.J. Wildl. Mgmt. 29(2): 39 7-398, Riney, T.1955.Evaluating condition of free-ranging red deer (Cervus elaphus), with special reference to New Zealand, New Zealand J. Sci. and Tech., Sec. B. 36(5): 429-463. Rush, W. M.1932.Bang's disease in the Yellowstone National Park buffalo and elk herds.J. Mammal, 13(4): 371 -372, Schwartz, J. E. and G. E. Mitchell.1945.The Roosevelt elk on the Olympic Peninsula, Washington,J. Wildi. Mgmt. 9(4): 295- 319. Simkin, D. W.1965.Reproduction and productivity of moose in Northwestern Ontario.J. Wildl, Mgmt. 29(4): 740-750,

67 Trainer, D. 0. and R. P. Hanson.1960.Leptospirosis and brucellosis serological reactors in Wisconsindeer, 1957-1958, J. Wildi. Mgmt. 24(1):44-5Z.

Verme, L. J.1965.Reproduction studies on penned white-tailed deer.J. Wildi. Mgmt. Z9(1): 74-79.

Verme, L. J.1967.Influence of experimental dietson white-tailed deer reproduction.Trans. N. Am. WildI. and Natur.Resour. Conf. 32:405-4Z0.

Welch, B. L.196Z.Adrenals of deer as indicators of populationcon- ditions for purposes of management.In:Proc. First National White-tailed Deer Dis. Sympo.pp. 94-108. Athens, Georgia, S. E. Sec. Wildi. Soc.ZOip.

. APPENDIX 1 Data resulting from examination of ovaries and uteri for Roosevelt elk collected in Western OregQn,1965, 1967and1968reproductive seasons.

Symbols: L and R = left and right uterine horn (location of embryo). cR/FR= crown-rump or forehead-rump length of embryos. r = regressed corpus luteum, Where more than one corpus luteum is indicated for a pregnant cow, the smaller presumably resulted from a postconception ovulation (Halazon and ,Buechner,1956). Table A.Ovarian and uterine data--pregnant Roosevelt elk collectedon Millicoma area, November 20 and 21, 1965.

Corporalutea Embryo Collection Diameterin mm Age Corpora Albicantia No. Follicle No. CR/FR (Left (Right) Uterine No. Date (Years) Udder ovary)ovary) 2mm 2-4mm 4-1-mm 2-5mm 5-8mm 8+mm Total horn mm Sex (Ovarian Exam. Complete)

65-is 11/20 4 is, 2 1 1. 4 2 0 6 70-80 * -31 11/20 4 - 15, 8 2 0 0 6 1 1(9mm) 8 - 68 M -33 11/20 4 16, 7 2 0 0 18 1 0 19 - 73 F -45 11/20 4 - - 17,8 1 0 0 11 3 0 14 60-70 M* -62 11/21 4 - 13,7 0 0 0 3 1 1(9mm) 5 R 105 F -25 11/20 5 15, 8 1 1 0 8 3 0 11 70-80 F* -74 11/21 5 14, 8 2 0 0 12 0 0 12 R 36 -27 11/20 6 - 1 14,7 1 0 26 3 0 29 L 80 M (Ovarian Exam. Incomplete)

-55 11/20 4 - 1 1 0 (left ovary only) R 25 - -61 11/21 4 - 17, 7 0 0 0 (right and part of left ovary) - 66 M

*(65-15, 25, & 45) Embryo damaged instorage. Table B. Ovarian and uterine data--pregnant Roosevelt elk collected on Millicomaarea. January 6-28, 1968

CorporaLutea Embryo Diameterin mm Collection Corpora Albicantia No. Follicle No. CR/FR Age (Left (Right Uterine No. Date (Years)Udder ovary) ovary)<2mm 2-4mm 4+mm 2-5mm 5-8mm 8+mm Total horn mm Sex (Ovarian Exam. Complete)

68-6 1/6 2 Wet 16,7 1 1 0 7 0 1(11mm) 8 192 F -4 1/7 3 Dry 16, - 1 0 0 17 0 0 17 168 M -7 1/6 3 Dry 15,5 0 0 0 5 2 1(8mm) 8 186 F -17 1/13 4 Dry 17,8 0 0 0 14 1 1(9mm) 16 L 289 M -22 1/14 4 Wet 16, 0 1 0 13 2 1(10mm) 16 - 174 M* -23 1/14 4 Wet 18,8 1 0 1 20 1 0 21 R 205 M -25 1/14 4 Dry - 14, 0 1 1 11 2 0 13 R 103 M -30 1/20 4 Dry 9 18, 0 0 1 21 3 0 24 R 284 M -45 1/28 4 Dry 16, 3 0 0 0 25 1 0 26 L 292 F -5 1/6 5 Dry 5 16, 0 0 0 8 1 1(8mm) 10 L 226 F 5 -14 1/7 Dry 18,7 7 1 0 24 2 2(8mm) 27 - 218 M -32 1/21 5 Dry 16, 2 0 0 12 2 0 14 - 232 F -13 1/7 6 Dry 18, - 4 0 0 36 1 0 37 L 177 M -29 1/20 6 Dry 7 18, 0 0 0 36 0 1(12mm) 37 R 139 F -34 1/21 6 Dry 19, 6 1 0 0 14 1 0 15 L 254 F -36 1/21 6 Dry 18, 4 0 0 0 31 2 1(9mm) 34 L 261 M -44 1/27 6 Dry 18,6 0 0 0 31 3 1(9mm) 35 R 335 F 67-1311/18 8 Wet 17,5 0 0 0 19 2 0 21 L 238 M 68-16 1/13 8 Wet 18,4 - 2 0 0 5 1 1(9mm) 7 213 M -24 1/14 8 Wet - 17,6 0 0 1 17 3 0 20 R 272 M 6 8 -37 1/27 Wet 20,3 2 0 1 11 1 0 12 R 202 F -8 1/6 9 Dry 6 13, 1 0 0 19 1 0 20 194 M 10 - -26 1/20 Wet 17, 0 0 0 14 1 0 15 R 236 F (Ovarian Exam. Incomplete)

-20 1/13 10 Dry -- 0 0 0 (right ovary only) 151 F *(68-22) Anterior end of mandible 5 mm shorter than opposing cranialsurface. Table C. Ovarian and uterine data--pregnant Roosevelt elk collected on Miflicoma area, November 2 - December 14, 1968. CorporaLutea Embryo Diameter in mm Corpora Albicantia No. FollicleNo. Collection CR/FR Age (Left (Right Uterine No. Date (Years)Udder ovary) ovary)<2mm 2-4mm 4+mm 2-Smm 5-8mm 8+mm Totalhorn mm Sex (Ovarian Exam. Complete)

68-85 11/3 2 Dry 15,8 0 0 0 14 1 0 15 R 30 -98 11/8 2 Dry 16,6 0 0 0 28 1 l(Smm) 30 L 35 -13912/14 2 Dry 14, 1 2 0 14 2 0 16 L 57 -79 11/2 3 Dry 12,4 0 0 0 14 1 l(llmm) 16 L 5 - -87 11/3 3 Dry 13,7 - 0 0 0 18 3 l(9mm) 22 R 50 -91 11/3 3 Dry 16, 0 0 0 5 2 l(lOmm) 8 L 54 -89 11/3 4 Dry 5 15, 0 0 0 6 1 l(llmm) 8 L 102 M -97 11/7 4 Wet 12, 6 0 2 0 14 2 0 16 L 15 -11211/10 4 Wet 10,3 0 1 0 17 1 0 18 R 11 -12512/14 4 Dry 14,7 6 1 0 0 9 1 l(Smm) 12 R 133 M l(lOmm) -94 11/5 S Dry 12,3 0 0 0 19 3 0 22 R 4 -96 11/6 5 Wet 12, 0 1 1 14 2 0 16 R 13 - -10211/9 5 Dry 15,7 0 0 0 25 4 l(9mm) 30 L 55 - -10511/9 5 Dry - 14,7 0 0 0 54 1 0 55 R 50 -10711/9 5 Wet 12,4 0 1 0 12 1 1(8mm) 14 - 6 -84 11/3 6 Wet 14,6 0 1 0 12 4 0 16 R 15 -99 11/8 6 Dry 13,6 1 0 0 29 0 2(9mm) 31 R 24 -10011/9 6 Wet - 14,6 1 0 0 16 1 l(Smm) 18 R 69 M -10111/9 6 Dry 1S,6 0 0 0 14 2 l(lOmm) 17 L 74 M -10411/9 6 Dry 16,7 0 0 0 9 2 l(lOmm) 12 L 70 F -11611/10 6 Wet 15,7 0 0 0 16 3 0 19 L 88 -82 11/3 7 Wet 11, 0 0 0 27 2 0 29 - -88 11/3 7 Dry 15,7 0 0 0 13 1 0 14 R 13 -92 11/4 7 Wet 14, 7 1 1 0 26 4 0 30 -11511/10 7 Dry 12, 41 3 0 0 37 1 l(lOmm) 39 L 15 -75 11/2 8 Wet 15, 8 3 1 0 12 1 l(9mm) 14 L 50 -77 11/2 8 Dry 14,6 0 0 0 26 3 0 29 L 53 -95 11/6 8 Wet 14,6 0 1 0 11 0 1(10mm) 12 L 33 -10611/9 8 Dry 14,8 0 0 0 23 1 l(Omm) 25 R 82 M -73 11/2 9 Wet 17, 0 1 0 8 5 0 13 L 39 -12612/14 9 Dry 15,9 1 0 0 31 3 0 34 L 44 (68-85, 112 & 84)Gelatinous exudate in cervix or vagina. /68_107) Chorion much fragmented; embryo appearing less developed than 6 mm embryos associated with unfragmented membranes. 4,(6882)Chorion much fragmented; embryo not located. ',(68-92)Embryo lost in field; chorion recovered from uterus. ' (68-115) The 4 mm corpus luteum appeared to be regressing; confirmed by histological examination. Table D. Ovarian and uterine data--pregnant Roosevelt elk collected on North Coastarea, November 11-18, 1967.

Corpora Lutea Embryo Diameter in mm Collection Corpora Albicantia No. Age (Left Follicle No. CR/FR (Right - Uterine No. Date (YearslUdder ovary) ovarr) 2mm 2-4mm 4+mm 2-5mm 5-8mm 8+mm Total horn mm Sex (Ovarian Exam. Complete)

67-72 11/11 2 Dry 15, 0 1 0 9 1 1(11mm) 11 L 4 3 -13911/18 14, 5 0 0 0 4 1 0 5 - 78 - -54 11/11 - 4 13, 5 0 1 0 22 3 0 25 - 86 F -79 11/11 4 Dry 8 17, 2 0 0 13 0 l(l5mm) 14 R 23 - -66 11/15 5 Dry 15,6 5 1 0 23 0 l(8mm) 24 58 -

-J Table E.Ovarian and uterine data--pregnant Roosevelt elk collected on North Coast area, November 16 and 17, 1968 and February 15, 1969.

CorporaLutea Embryo Diameter in mm Collection Corpora Albicantia No. Follicle No. CR/FR Age (Left (Right Uterine No. Date (Years)Udder ovary) ovary)<2mm 2-4mm 4+mm 2-5mm 5-8mm 84-mm Total horn mm Sex (Ovarian Exam.Complete)

68-156 2/15 1 Dry 11, 0 0 0 26 0 0 26 L 192 F -16311/17 2 Dry 14, 0 1 0 13 2 1(9mm) 16 L 3 -17211/17 4 Dry 13, 1 1 0 30 1 0 31 R 27 - -20811/16 4 7 14, 0 1 0 8 1 l(9mm) 10 R 26 - -20911/16 6 Wet - 16, 0 1 0 22 1 1(9mm) 24 - - -* (Ovarian Exams Incomplete)

-18011/16 5 Dry - -- - (both ovaries cut off) R 27

*(68-209) Embryolostin field, thorion recovered from uterus.

-J Table F. Ovarian and uterine data--pregnant Roosevelt elkcollected on Loon Lake area, November 17December 15, 1968.

Corpora Lutea Embryo Collection Diameter in mm Corpora Albicantia No. FollicleNo. Age (Left (Right UterineCR No. Date (Year) Udder ovary) ovary) <2 mm 2-4mm 4+mm 2-5mm 5-8mm 8+mm Total horn mm Sex (Ovarian Exam. Complete)

68-14812/14 1 Dry 15,5 0 0 0 16 1 1(9mm) 18 R 72 F -13312/7 2 Dry 14,7 0 0 0 19 1 1(10mm) 21 L 83 M -12111/17 4 Dry 16,10 0 0 0 19 2 0 21 L 75 M -12912/7 4 Wet 15,9 2 1 0 14 1 1(9mm) 17 R 110 F 1(12mm) -13612/7 4 Dry 14, 8 0 1 0 6 1 2(10mm) 9 R 115 F -14712/14 4 Wet 12, 9 0 2 0 19 1 1(8mm) 21 L 32 - -13412/7 5 Dry 13, 3 1 0 18 3 1(11mm) 22 L 4 - -13512/7 S Dry 14,8 2 0 0 20 4 0 24 L 107 -15112/15 5 Wet M 15,5 1 1 0 15 2 0 17 -127 R 120 F 12/7 7 Wet 8 17, 0 0 1 20 1 1(11mm) 22 R 101 M -14312/14 7 Wet 16,5 1 1 0 11 1 1(10mm) 13 - 116 F

-J Table G. Ovarian and uterine data- -nonpregnant Roosevelt elk collectedon Millicoma area, November 20 and 21, 1965. * CorporaLutea Collection Diameter in mm Age (Left (Right Corora Albicantia No. FollicleNo. No. Date (Years) Udder ovary) ovary) <2mm 2-4mm 4+mm 2-5mm 5-8mm 8+mm Total (Ovarian Exam. Complete)

65-17 11/20 1 - - - 0 0 0 16 0 l(lOmm) 17 -66 11/21 1 - - - 0 0 0 12 2 0 14 -67 11/21 1 - - - 0 0 0 21 1 0 22 -8 11/20 2 - - 0 0 0 30 0 l(9mm) 31 -46 11/20 2 - - - 0 0 0 27 1 1(l0mm) 29 -70 11/21 2 - - - 2 0 0 22 3 0 25 -14 11/20 3 - 1 0 1 16 0 1(9mm) 17 -63 11/21 3 - - - 0 0 0 15 0 l(9mm) 16 -71 11/21 3 7r 2 0 0 16 0 l(lOmm) 17 -12 11/20 4 - - 1 1 1 28 2 0 30 -18 11/20 4 - 4r 0 1 1 17 0 0 17 -73 11/21 4 - 12 - 0 0 1 14 1 0 15 -24 11/20 5 - - 7r 1 2 2 15 1 0 16 -58 11/20 5 - 0 1 1 15 1 0 16 -72 11/21 5 - - - 1 0 0 4 2 l(9mm) 7 -4 11/20 6 - - - 0 1 1 21 1 0 22 -6 11/20 6 - 6r 1 0 1 12 3 0 15 -51 11/20 6 - 6r 1 1 1 4 2 0 6 -9 11/20 7 - 0 0 0 15 1 0 16 -20 11/20 7 - - - 1 3 0 0 0 1(9mm) 1 -64 11/21 7 13 7r 0 0 2 22 1 0 23 -22 11/20 Aged - Adult 0 0 0 2 2 0 4 -35 11/20 Aged Adult - 0 0 1 13 1 0 14 (Ovarian Exam. Incomplete)

-30 11/20 2 0 0 0 (right ovary only) -10 11/20 3 8 r 5 r 0 0 1 (right&part of left ovary) -36 11/20 3 - - 1 0 1 (left ovary only) -5 11/20 4 - - 0 0 1 ( -7 11/20 4 - - - 0 0 0 (right ovary only) *Uteri (nongravid) from two yearlings, two two year olds,two three year olds, one four year old, and one five year old were examined forpregnancy at checking station, but are excluded from table since the specimenswere not available for reexamination. Table H. Ovarian and uterine data--nonpregnant Roosevelt elk collected on Millicomaarea, January 3February 25, 1968. CorporaLutea Collection ::eter Age (Right Corpora AlbicantiaNo. FollicleNo. No. Date (Years) Udder ovary' ovary) <2mm 2-4mm 4+mm 2-5mm 5-8mm 8+mm Total (Ovarian Exam. Complete)

68-57 2/25 1 Dry - 0 0 0 37 1 0 38 -43 1/28 2 Dry - - 0 0 0 22 2 0 -2 1/3 3 24 Dry - 0 0 0 -18 17 1 0 18 1/13 3 Dry - - 1 3 0 21 1 0 -40 1/27 3 Wet - - 221 0 1 0 -1 22 2 0 1/3 4 Dry - 1 0 0 32 0 1(14mm) 33 -21 1/14 4 Wet - 0 1 2 28 0 0 -33 1/21 5 28 Dry - - 3 0 0 -35 11 1 1(10mm) 13 1/21 5 Dry 12, - 3 2 0 4 2 -3 1/6 6 1(9mm) 7 Wet - 0 1 1 26 1 0 27 -41 1/27 6 Wet - - 0 0 0 14 0 0 14 -9 1/6 7 Wet - 0 0 1 12 1 0 -15 1/7 7 Wet 13j' 1 1 1 26 0 0 67-132 1/10 8 Dry - 12, 2 0 0 24 0 2' 6r 1(9mm) 2.5 68-11 1/7 8 Dry 0 0 0 12 1 -12 1/13 8 1(12mm) 14 Wet - 2 1 0 7 1 0 -19 1/19 9 Wet 0 1 0 30 0 0 30 -31 1/20 9 Wet - - 0 1 0 20 0 67-130 1/2.5 0 11-15 Dry - - 0 0 0 68-42 1/27 1 0 0 1 11-15 Wet - - 0 2 0 23 1 0 24 -58 2/25 11-15 - Dry - 0 0 0 -38 1/28 0 0 0 0 15+ Dry - 0 0 0 2 0 0 2 (Ovarian Exam. Incomplete)

-10 1/7 4 Dry - 7, 0 1 0 V -27 (right ovary only) 1/20 5 Dry - - 3 1 0 (" -28 1/21 7 " ) Wet - 0 1 0 (" " ") 68-40, 9, 15, 19, 31 & 28)Gelatinous exudate in cervixor vagina. (68-10)Corpus luteum forming in ruptured follicle. Table I.Ovarian and uterine data--nonpregnant Roosevelt elk collected on Millicomaarea, November 2December 14, 1968. CorporaLutea Diameter in mm Collection Corpora Albicantia Age (Left (Right No. FollicleNo. No. Date (Years) Udder ovary) ovary) <2mm 2-4mm 4+mm 2-Smm 5-8mm 8+mm Total (OvarianExam. Complete) 68-76 11/2 1 1 Dry - 0 0 0 10 2 0 12' -83 1 - 11/3 Dry 0 0 0 21 2 1(8mm) 24 -103 11/9 1 Dry - 0 0 0 40 0 1(9mm) 4111 -111 1 11/10 Dry 0 0 0 20 3 0 23 -74 11/2 2 Dry - 5 r 0 0 0 10 4 1(11mm) 15 -78 11/2 2 Dry - 0 0 0 20 2 1(8mm) 232, -86 2 11/4 Dry 11 r 1 0 0 11 2 1(9mm) 14' 4r -110 11/10 2 Dry . - 0 0 0 18 2 0 20 -80 11/2 3 Wet - - 0 1 0 13 0 0 1311 -108 11/9 4 Wet - 0 1 0 33 2 0 35 -109 11/10 4 Wet 0 0 1 8 0 1(10mm) 9'4 -118 11/10 4 Wet - 13,4 r 1 1 0 18 0 1(9mm) 19 -120 11/16 4 Wet 0 1 0 14 2 0 164 -81 5 - 11/2 Wet 11, 4r 0 0 1 7 1 1(11mm) -90 11/3 5 Dry - 2 0 0 17 3 0 20 -93 11/5 5 Wet 6r 4r 0 1 0 12 4 1(12mm) 17 -117 5 11/10 Dry 6 r 15, 1 0 0 13 1 1(9mm) l5 -122 11/18 6 Wet 4r 0 0 0 4 0 0 4' 8 -72 11/2 Wet - 0 0 0 9 1 0 10 -113 11/10 9 Wet 6 r 1 1 0 9 0 1(9mm) 10k, -123 11/22 11-15 Wet - 0 0 0 6 0 0 6' -124 12/14 11-15 Wet 0 1 0 4 2 0 6 (Ovarian Exam. Incomplete) -114 11/10 8 Dry - - 1 0 0 11 2 0(left ovary only; right cystic) J (68-76, 103, 80, 109, 122 & 123)Gelatinous exudate in cervix or vagina. /68-86) Corpora lutea had macroscopic appearance of being functional, but histological examination indicated bothwere regressing. (68-118)Embryonic mortality--fragments of disintegrating chorion present in tips of both uterine horns; histological examinationshowed necrotic ,,tissue structure; embryo not located. ,(68_118 & 81)Indicated status of corpora lutea verified by histological study. (68-114)Right ovary contained two cystic follicles, 40 mm x 40 mm, and 4 mm x 20 mm, respectively. Table J.Ovarian and uterine data-nonpregnant Roosevelt elk collectedon North Coast area, November 11 - December 6, 1967. Corpora Lutea Diameter in mm Collection CorporaAlbicantiaNo. Age (Left (Right Follicle No. No. Date (Years) Udder ovary) ovary) <.2mm 2-4mm 4+mm 2-5mm 5-8mm 8+mm Total (Ovarian Exam. Complete)

67-137 - 12/6 calf Dry 0 0 0 22 1 0 23 -56 11/11 1 Dry - 0 0 0 10 1 0 11 -138 11/16 1 Dry 0 0 0 18 2 0 20 -58 11/12 2 - - Dry 0 0 0 24 1 1(11mm) -62 11/11 3 - Dry lOr 0 0 1 3 1 1(9mm) 5 -77 11/11 3 Wet 0 1 1 27 1 0 28 -93 11/11 3 4r 12, 2 0 0 24 0 1(10mm) 25 -128 11/11 11-15 Wet 12, 1 1 0 48 0 0 48 (Ovarian Exam. Incomplete)

-92 11/18 4 - - - 0 0 1 (right ovary only)2 -141 11/18 5 - - - 0 0 1 (left ovary only)

'(67-62& 93)Suspected embryonic mortality--uterus contained membrane fragments (1mm to 3 mm) that histologically resembled chorionic tissue.

'(67-l41)Embryonic mortality--densely compacted tissue mass 12mm x 6 mm x 3 mm present in tip of right uterine horn; histological examination revealed many nucleated red blood cells that resembled embryonic red bloodcells of healthy thorion. Table K. Ovarian and uterine data--rionpregnant Roosevelt elk collected on North Coastarea, November 16 and 17, 1968.

Corpora Lutea Collection Diameter in mm Age (Left (Right Corpora AlbicantiaNo. Follicle No. No. Date (Years) Udder ovary) ovary) (2mm 2-4mm 4+mm 2-5mm 5-8mm 8+mm Total (Ovarian Exam. Complete)

68-207 11/16 1 Dry 0 0 0 -210 9 2 0 11 11/17 1 Dry - 0 0 0 20 0 1(9mm) 21 -174 11/16 2 Dry 11, - 0 1 0 19 1 0 -181 11/16 4 Wet 20 10, 1 1 0 -173 7 1 1(9mm) 9 11/17 8 - - - 0 0 1 3 0 1(l0mm) 4

0 Table L.Ovarian and uterine data--nonpregnant Roosevelt elkcollected on Loon Lake area, December 7-15, 1968. Corpora Lutea Collection Diameter in mm Age (Left (Right Corpora AlbicantiaNo. FollicleNo. No. Date (Years) Udder ovary) ovary) (2mm 2-4mm 4+mni 2-5mm 5-8mm 8+mm Total (Ovarian Exam. Complete)

68-144 12/14 calf Dry 0 0 0 0 0 0 0 -146 12/14 1 Dry 0 0 0 14 1 l(l0mm) -149 12/15 1 Dry l6ti 0 0 0 9 1 0 10 -142 12/14 2 Dry 0 1 0 7 0 1(8mm) 8 -131 12/7 3 Dry 6r 2 1 0 27 0 1(10mm) 28 -145 12/14 3 5 r 3 1 0 12 1 l(8mm) 14,, -130 12/7 4 Dry Sr 0 2 0 21 0 4r 1(12mm) 22 -152 12/15 4 Wet - 11 2 0 1 0 37 1 1(9mm) 39 4r -138 12/7 5 Wet 6 r 1 1 1 9 1 1(8mm) 11 -141 12/14 5 Wet 4 1 0 30 2 0 32 -137 12/8 6 Wet 3 3 0 28 3 0 31 -128 1217 7 Wet 4r 2 2 1 19 0 1(12mm) 20 -150 12/15 7 Wet 1 1 0 16 1 0 173 -132 12/7 11-15 Wet 13, 3 2 0 10 1 2(9mm) l3 L'(68_149)Gelatinous exudate in cervix or vagina.

30 & 152)Indicated status of corpora lutea verified by histologicalexamination. '(68-132) Embryonic mortality--two pieces of disintegratingchorion (8 mm x 15 mm & 4 mm x 11 mm)present in uterus; histological study showed tissue to be necrotic, and that corpus luteumwas apparently functional. E31 APPENDIX Z Data resulting from examination of ovaries and uteri for Rocky Mountain elk collected in Northeast Oregon,1967and1968reproduc- tive Seasons.

Symbols: L and R = left and right uterine horn (location of embryo). CR/FR= crown-rump or forehead-rump length of embryos. r = regressed corpus luteum. Where more than one corpus luteum is indicated for a pregnant cow, the smaller presumably resulted from a postconception ovulation (Halazon and Buechner,1956). Table A. Ovarian and uterine data--pregnant Rocky Mountain elk collected in Northeast Oregon, November ii, 1967january 20, 1968. CorporaLutea Embryo Collection Diameter in mm Age (Left Corpora AlbicantiaNo. FollicleNo. CR/FR (Right Uterine No. Date (Years) Udder ovary)ovary) <2mm 2-4mm4+mm 2-5mm5-8mm 81-mm Total horn mm Sex (Ovarian Exam. Complete) 67-213il/il 2 Wet 13, 7 0 0 0 37 0 l(lOmm) 38 L 29 -22211/12 2 Dry 7 13, 2 0 0 19 0 1(9mm) 20 R 29 -22311/11 2 Dry 13,10 0 0 0 73 1 1(9mm) 73 R 27 7 -24311/12 2 Dry 13, 7 0 0 0 41 1 0 42 L 17 -13 3 12/22 Dry 18, 10 2 1 1 11 1 0 12 L 149 F -14 3 12/21 Wet 15,9 1 0 1 15 0 1(9mm) 16 R 116 F -17011/11 3 Wet 14, 7 0 2 0 44 0 1(8mm) 45 - 16 -17211/11 3 Wet 15,7 0 1 0 20 0 1(8mm) -211 21 L 6 11/11 3 Wet 8 13, 0 1 1 22 1 0 23 R 26 -22011/11 3 Wet 15, 8 0 1 0 27 1 1(9mm) -221 29 L 36 11/il 3 Wet 16, 1 0 0 14 1 0 15 R 12 -22411/11 3 Wet 13, 9 0 0 1 22 2 0 24 R 13 -23111/12 3 Dry 13, 0 0 0 10 1 1(8mm) 12 L 11 - -24011/12 3 Wet 13, 8 0 2 0 16 1 1(9mm) 18 L 15 -248 1/20 3 Dry 10, 16, 0 0 0 20 1 1(9mm) 22 R 245 F -17311/11 4 - - 14,7 0 1 1 41 0 1(8mm) 42 R 21 -21211/12 4 Wet 10 14, 1 2 0 29 3 0 32 L 21 -21611/19 4 Wet 14, 6 2 3 0 19 0 1(9mm) 20 L 18 -22511/11 4 Wet 13,4 0 2 0 13 1 0 14 R 13 -24712/Il 4 Wet 13,7 2 0 1 21 3 1(9mm) 25 L 32 -23411/12 5 Wet 14, 6 1 0 0 17 1 1(8mm) 19 L 37 - -24211/11 S Wet 14,7 0 0 0 16 2 0 18 R 18 - -22911/12 6 Wet 13,7 0 1 1 23 2 0 25 11/11 R 19 -233 6 Dry 6 15, 0 1 0 19 3 0 22 R 26 -21511/19 7 Wet 14,6 0 3 0 27 2 2(8mm) 31 R 23 -218 8 11/19 8 16, 0 0 1 25 0 1(9mm) 26 R 75 F -23011/12 8 Wet 8 14, 1 1 0 38 2 0 40 L 20 -21411/13 9 5 13, 0 1 0 23 1 0 24 L 17 -22811/12 9 Dry 7 13, 2 0 0 44 3 0 -189 47 R 25 11/13 11-15 Dry 12, 0 1 1 6 0 1(9mm) 7 R 7 (Ovarian Exam, Incomplete)

-16411/18 3 Wet 13,5 r 0 0 1 (left ovary only) L 6 -25812/16 4 Wet - -- (gravid uterus, lost after cell.) - - -68 1/19 4 - - ( ' " -66 1/16 5 8 - 0 0 0 (left ovary only)"' ) R 111 M -67 1/16 5 - - (embryo oniy collected) - -232 10 97 F 11/12 Dry 12, 7 1 1 0 (right & part of left ovary) L 15 - PD Table B.Ovarian and uterine data--pregnant Rocky Mountain elk collected inNortheast Oregon, November 16, 1968 -January 25, 1969.

Corpora Lutea Embryo Diameter in mm Collection Corpora Albicantia No. FollicleNo. Age (Left (Right UterineCR/FR No. Date (Years) Udder ovary) ovary) <2mm 2-4mm4+mm 2-Smm5-8mm 8+mm Total horn mm Sex (Ovarian Exam. Complete) 67-17811/16 1 Dry 13, 6 0 0 0 16 2 l(9mm) 19 R 35 - 68-23211/16 1 Dry 11, 4 0 0 0 10 2 0 12 L 15 - 68-31811/17 1 Dry 13, 4 0 0 0 16 1 0 17 L 19 - 68-33011/18 1 Dry 13, 7 0 0 0 26 1 0 27 R 30 - 67-27512/22 2 Dry 13,6 0 0 0 13 1 0 14 L 139 M 67-28012/7 2 Dry 12,5 0 0 0 27 1 0 28 R 114 F 68-26711/19 2 Wet 12, 0 1 0 34 2 0 36 L 28 68-28011/16 2 6 14, 0 1 0 9 1 0 10 R 61 68-30312/20 2 14, 4 0 0 0 5 0 1(8mm) 6 L 114 M 68-33211/17 2 Dry 12, 0 0 0 16 2 0 18 R 24 - 67-27312/22 3 Wet 11,5 0 1 0 22 0 0 22 L 101 M 67-28112/8 3 Dry 7 15, 0 0 0 5 1 0 6 L 105 M 68-268 11/18 3 Wet 13, 1 1 0 1 0 l(lOmm) 2 R 16 68-28111/16 3 Wet 6 15, 1 1 0 22 1 l(8mm) 24 R 10 68-28411/18 3 Wet 13,7 0 2 0 19 2 l(lOmm) 22 - 30 - 68-336 11/18 3 Wet 1 14,7 0 1 18 1 0 19 L 24 - 67-18411/16 4 Wet 5 15, 0 0 0 20 3 0 23 L 58 - 67-27712/8 4 Wet 1 13,6 0 0 17 2 0 19 R 103 F 67-28212/7 4 Dry 7 15, 0 1 0 6 2 0 8 R 144 F2, 68-23311/17 4 Wet 10 15, 0 1 0 28 2 1(8mm) 31 - - 68-23412/8 4 Wet 13,9 1 1 0 4 1 0 5 R 82 M 68-27911/16 4 Wet 1 17,7 0 0 20 2 0 22 L 26 - 68-28611/16 4 Wet 15,8 0 1 1 36 2 0 38 L 47 - 68-287 11/16 4 Wet 12, 0 0 1 18 2 0 20 5 - 68-28911/16 4 Wet - 15,6 0 1 0 15 1 1(8mm) 17 R 66 M 68-32311/20 4 Wet 7 16, 0 1 0 4 0 1(8mm) 5 L 28 68-32411/20 4 Wet 15,7 0 2 0 21 0 l(lOmm) 22 L 26 68-31711/17 5 Wet 14,3 0 0 1 8 3 0 11 R 51 - Continued on next page Table B continued.

Corpora Lutea Embryo Collection Diameter in mm Age Corpora Albicantia No. Follicle No. CR/FR (Left (Right Uterine No. Date (Years) Udder ova) ovary) <2mm 2-4mm4+mm 2-5mm S-8mm 8+mm Total horn mm Sex (Ovarian Exam.Completecont.)

68-33711/16 5 Wet 15, 0 1 0 4 1 0 5 L 32 - 67-27912/8 6 Dry 15, 7 1 1 0 18 1 1(8mm) 20 L 114 F 68-32911/19 6 Wet 13,9 1 2 0 13 2 1(9mm) 16 R 28 - 68-29011/17 7 Wet 7 14, 4 0 0 18 0 2(9mm) 20 R 83 M 67-18111/16 8 - 13,5 1 1 0 25 2 0 27 R 35 - 67-27112/31 8 Wet 11,6 1 0 0 7 1 0 8 L 179 M 68-23011/16 8 Wet 13, 0 0 0 11 1 0 12 4 - 68-33111/16 8 Wet 7 11, 1 1 0 7 1 l(8mm) 9 L 72 M 67-18611/16 9 15, 7 0 0 0 25 1 0 26 L 89 F 68-32711/16 9 Wet 12, 0 1 0 1 2 0 3 L 12 - 68-33511/16 9 Wet 4 15, 1 4 0 6 1 0 7 R 61 (Ovarian Exam. Incomplete) 68-31611/17 1 Dry - 0 0 0 (right ovary only) L 16 - 67-18511/16 2 Wet - - - - - (ovaries cut off) L 15 - 68-28311/16 2 - - 0 0 0 (right ovary only) R 25 - 67-18311/16 3 Dry - 1 0 0 ( " " " ) L 45 - 67-17611/16 4 Wet 14, 7 0 1 0 (left ovary only) L 22 - 68-69 1/21 5 Wet - - - - - (embryo only collected) - 228 M 68-32811/19 5 Wet - 1 1 0 (right ovary only) - 95 F 67-27412/26 9 Wet 13,6 0 0 0 - (left ovary only) 196 M 67-257 1/2511-15 Wet - - - - - (embryo only collected) - 208 F !J(68_281) Embryo apparently alive, but lacking appendages; classifiedas a teratism. L'(68233) Embryo lost in field; chorion recovered fromuterus. Table C. Ovarian and uterine data--nonpregnant Rocky Mountain elkcollected in Northeast Oregon, November 11-19, 1967.

Corpora Lutea Diameter in mm Collection Corpora Age (Left (Right) AlbicantiaNo. Follicle No. No. Date (Years) Udder ovary) ovary) <2mm 2-4mm 4+mm 2-5mm 5-8mm 8+mm Total (Ovarian Exam. Complete)

67-217 11/19 calf Dry - 0 0 0 3 1 0 4 -219 11/11 calf - - Dry 0 0 0 20 0 0 20 -187 11/11 1 Dry - 0 0 0 21 2 0 23 -188 11/11 1 Dry - 0 0 0 15 1 0 16 -218 11/19 1 Dry 0 0 0 21 1 0 22 -227 11/12 1 Dry - - 0 0 0 25 0 0 25 -171 11/12 3 Wet - - 0 0 1 30 1 1(8mm) 32 -241 11/11 4 Wet 4r 0 0 1 41 1 0 42 -169 11/11 8 Wet 13, 5r 1 0 1 9 2 1(8mm) 12 -226 11/11 10 Wet 12, 0 2 0 12 2 1b2mm) 15 (Ovarian Exam. Incomplete)

-168 11/13 1 Dry 0 0 0 (left ovary only) -190 11/11 3 Wet 13, - 0 0 0 ("" " )*

*(67-190) Embryonic mortality suspected--uteruscontained membrane fragments (1 mm3 mm) that histologically resembled chorionic tissue. Table D. Ovarian and uterinedata--nonpregnant Rocky Mountain elk collected in Northeast Oregon, November 16and 17, and December 22, 1968. Corpora Lutea Collection Diameter in mm Age (Left (Right CorporaAlbicantiaNo. Follicle No. No. Date (Years) Udder ovary) ovary) <2mm 2-4mm 4+rnm 2-5mm 5-8mm 8+mm Total (Ovarian Exam. Complete) 68-281 11/17 1 Dry 4r 0 0 0 26 1 68-285 11/17 1 0 27 Dry - 0 0 0 19 1 68-326 11/17 1 Dry 0 20 Sr 0 0 0 12 1 68-334 11/17 1 0 13 Dry - 0 0 67-180 0 23 1 0 24 11/16 2 Dry 6r - 0 0 0 17 3 0 67-272 12/22 3 Wet 11. 20 0 0 1 25 1 68-333 11/16 3 0 26 Wet 14, 1 2 68-228 11/16 1 19 1 0 4 Wet 13, 1 1 0 24 1 68-315 11/16 4 1(8mm) 26 Wet 16, 1 0 1 10 1 1(9mm) 12 (Ovarian Exam. Incomplete)

68-338 11/16 1 Dry (ovaries cut off) 67-179 11/16 10 Wet 8r 2 2/ 0 1 9 0 1(lOmm)'

'(68-333)Embryonic mortality--fragments of disintegrating chorion (largest 7mm x 18 mm) present in tips of uterine horns; tion indicated necrotic tissuestructure; embryo not located. histological examina- "(67-179)Right ovary only. I

Appendix Table 3. Sex of Roosevel.t and RockyMountain elk fetuses collectedin Oregon, 1965, 1967 and 1968 reproductiveseasons* Total Fetuses Males per Males Females Sexed 100 Females Percent Males Roosevelt elk 2.7 22. 49 12.3 55 Rocky Mountain elk 12 12 24 100 50

*Sex determined only forfetuses with forehead-rumplength of at least 65mm. Appendix T.abie 4. Data on physical condition for pregnant Roosevelt elk collectedon Millicoma area, 1967 reproductive season.

Collection Embryo Percent Fat Body Weights Total Kidney Metatarsal Weight of CR/FR (Pounds) cg Adrenal per No. Date Age Fat Marrow Adrenals lb Hog-dressed 68- (1968) (Years) Udder (mm) Sex Index (Wet Basis) Whole Hog-dressed (gL Body Weight

6 1/6 2 Wet 192 F 27 79 480 324 6.70 2.07 4 1/7 3 Dry 168 M 95 89 526 354 4.97 140 7 1/6 3 Dry 186 F 127 87 526 365 4.35 1.19 17 1/13 4 Dry 289 M 43 90 596 428 5.23 1.22 22 1/14 4 Wet 174 M 32 89 623 418 6.01 1.44 23 1/14 4 Wet 205 M 33 90 546 366 5.66 1.,5 25 1/14 4 Dry 103 M 61 89 578 402 5.08 L26 30 1/20 4 Dry 284 M 80 89 576 404 6.08 1.50 45 1/28 4 Dry 292 F 57 88 577 409 4.85 1.19 5 1/6 5 Dry 226 F 64 74 658 428 4.60 1.07 14 1/7 5 Dry 218 M 111 91 69& 462 5.41 L17 32 1/21 5 Dry 232 F 43 89 633 420 6.43 1.53 13 1/7 6 Dry 177 M 85 88 644 432 6.27 1.45 29 1/20 6 Dry 139 F 63 88 606 415 6.26 1.51 34 1/21 6 Dry 254 F 52 90 612 417 5.67 1.36 36 1/21 6 Dry 261 M 49 89 572 405 4.92 1.21 44 1/27 6 Dry 335 F 104 88 619 425 6.33 1.49 24 1/14 8 Wet 272 M 32 86 626 401 6.12 1.53 37 1/27 8 Wet 202 F 14 33 569 366 6.91 1.89 8 1/6 9 Dry 194 M 69 82 647 435 7.33 1.69 26 1/20 10 Wet 236 F 27 81 614 379 8.01 2.11 Appendix Table 5. Data on physical condition for nonpregnant Roosevelt elk collectedon Millicorna area, 1967 reproductive season. Collection Corpus Percent Fat Body Weights Total Luteum Kidney Metatarsal Weight of cg Adrenal per No. Date (Pounds) Age Present Fat Marrow Adrenals lb Hog-dressed 68- (1968) (Years) Udder (x) Index (Wet Basis) Whole Hog-dressed (g) Body Weight

57 2/25 1 Dry 16 70 432 43 1/28 2 Dry 60 91 495 351 3.61 1.03 2 1/3 3 Dry 30 84 537 367 5.62 1.53 18 1/13 3 Dry 31 86 524 395 4.42 1.12 40 1/27 3 Wet 13 38 529 356 4.24 1.19 1 4 1/3 Dry 30 89 - 5.73 10 1/7 4 Dry x 32 88 628 446 5.30 1.19 21 1/14 4 Wet 40 82 614 390 5.24 1.34 27 1/20 5 Dry 67 90 586 408 6.64 1.63 33 1/21 5 Dry 23 90 582 415 3.98 0.96 35 1/21 5 Dry x 50 90 586 408 4.72 .1.15 3 1/6 6 Wet 18 68 602 371 5.86 1.58 41 1/27 6 Wet 7 57 591 360 5.65 1.57 9 1/6 7 Wet 12 46 596 408 6.79 1.66 15 1/7 7 Wet 14 43 600 408 5.54 1.36 28 1/21 7 Wet 10 11 601 394 6.31 1.60 11 1/7 8 Dry 32 88 658 444 5.59 1.26 12 1/13 8 Wet 9 20 562 371 6.98 ,l.88 19 1/19 9 Wet 5 12 577 376 6.06 1.61 31 1/20 9 Wet 20 65 568 356 7.64 2.15 42 1/27 11-15 Wet 15 67 581 378 6.61 1.75 58 2/25 11-15 Dry 6 1 6.20 38 1/28 15+ Dry 16 21 519 316 6.17 1.95 Appendix Tab].e 6.Data on physical condition for pregnant Roosevelt elk collected in Western Oregon, 1968reproductive season. Collection Embryo Percent Fat Body Weight Kidney Metatarsal Weight of CR/FR (Pounds) cg Adrenal per Area Date Age Fat Marrow Adrenals lb Hog-dressed 68- No. (1968) (Years) Udder (mm) Sex Index (Wet Basis) Whole Hog-dressed (g) Body Weight

C 148 12/14 1 Dry 72 F 96 89 B 85 11/3 2 Dry 30 - 101 88 503 360 4.00 1.11 B 98 11/8 2 Dry 35 198 91 543 391 4.21 1.08 C 133 12/7 2 Dry 83 M 175 89 - A 163 11/17 2 Dry 3 156 - - - B 79 11/2 3 Dry 5 - 137 90 527 388 3.82 0.98 B 87 11/3 3 Dry 50 256 89 586 413 6.00 1.45 B 91 11/3 3 Dry 54 266 90 600 427 5.65 1.32 B 89 11/3 4 Dry 102 M 163 90 552 429 4.07 0.95 B 97 11/7 4 Wet 15 100 91 661 455 5.22 1.15 B 112 11/10 4 Wet 11 36 86 491 334 5.01 1.50 121 C 11/17 4 Dry 75 M 114 89 - B 125 12/14 4 Dry 133 M 65 89 593 364 3.81 1.05 C 136 12/7 4 Dry 115 F 137 87 A 172 11/17 4 Dry 27 164 91 B 94 11/5 5 Dry 4 - 102 91 591 421 5.37 1.28 B 96 11/6 5 Wet 13 48 88 563 397 4.82 1.21 B 102 11/9 5 Dry 55 - 232 91 674 492 6.51 1.32 B 105 11/9 5 Dry 50 187 90 601 420 5.04 1.20 B 107 11/9 5 Wet 6 - 62 91 581 410 5.72 1.40 C 134 12/7 5 Dry 4 - 165 89 - - 135 C 12/7 5 Dry 107 M 228 88 - C 151 12/15 5 Wet 120 F 44 89 B 84 11/3 6 Wet 15 80 90 652 455 5.73 1.26 B 99 11/8 6 Dry 24 141 91 675 490 6.41 1.31 B 100 11/9 6 Wet 69 M 63 92 605 411 6.17 1.50 B 101 11/9 6 Dry 74 M 284 91 599 435 4.70 1.08 B 104 11/9 6 Dry 70 F 218 90 630 455 5.90 1.30 Continued on next page Appendix Table 6 Continued. Collection Embryo Percent Fat Body Weight Total Kidney Metatarsal Weight of cg Adrenal per Area Date Age CR/FR (Pounds) Fat Marrow ______Adrenals lb Hog-dressed 68- No. (1968) (Years) Udder (mm) Sex Index (Wet Basis) Whole Hog-dressed (g) Body Weight B 116 11/10 6 Wet 88 M 148 88 521 375 5.06 1.35 A 209 11/16 6 Wet 122 90 - - - B 82 11/3 7 Wet 73 91 595 405 4.96 1.22 B 88 11/3 7 Dry 13 - 190 88 639 439 4.88 (.11 B 92 11/4 - 7 Wet 100 92 646 435 5.74 1.32 B 115 11/10 7 Dry 15 180 89 671 499 4.43 0.89 C 127 12/7 7 Wet 101 M 50 88 - - B 75 11/2 8 Wet 50 - 61 90 631 477 4.58 0.96 B 77 11/2 8 Dry 53 - 175 92 619 440 6.62 1.50 B 95 11/6 8 Wet 33 - 32 88 591 397 5.90 1.49 B 106 11/9 8 Dry 82 M 194 90 631 435 5.62 1.29 B 73 11/2 9 Wet 39 - 73 91. 729 495 5.93 1.20 B 126 12/14 9 Dry 44 - 48 90 643 424 7.20 1.70

A = North Coast area. B Millicoma area C Loon Lake area VEmbryonot measured. 'Chorionfragmented; embryo not located. Appendix Table 7. Data on iihysical condition for nonpregnant Roosevelt elkcollected in Western Oregon, 1968 reproductiveseason. Collection Percent Fat Body Weights Total Ovarian! Kidney Metatarsal Weight of cg Adrenal per Area Date Age (Pounds) Uterine Fat Marrow Adrenals lb Hog-dressed 68- No. (1968) (Years) Udder Data Index (Wet Basis) Whole Hog-dressed (g) Body Weight

B 76 11/2 1 Dry 60 89 443 319 4.76 1.49 B 83 11/3 1 Dry 123 90 481 344 3.77 1.10 B 103 11/9 1 Dry 49 87 465 319 4.19 1.31 B 111 11/10 1 Dry 35 86 453 311 3.86 1.24 C 149 12/15 1 Dry 19 88 A 207 11/16 1 Dry 94 - - - A 210 1 11/17 Dry 66 - - B 74 11/2 2 Dry 119 - 549 380 4.97 1.31 B 78 11/2 2 Dry 73 90 535 389 4.71 1.21 B 86 11/4 2 Dry V 115 90 539 379 3.81 1.01 B 110 11/10 2 Dry 104 90 561 392 3.31 0.84 A 174 2 11/16 Dry 140 88 - B 80 11/2 3 Wet 13 36 561 385 5.29 1.37 B 108 11/9 4 Wet 28 89 571 399 4.05 1.02 B 109 11/10 4 Wet 22 89 551 389 5.14 1.32 B 118 11/10 4 Wet /,3j 64 88 586 393 4.51 1.15 B 120 11/16 4 Wet 47 85 601 420 5.97 1.42 C 152 12/15 4 Wet V 37 87 - A 181 11/16 4 Wet V 56 85 B 81 11/2 5 Wet V 30 87 663 473 6.60 1.40 B 90 11/3 5 Dry 50 88 519 354 6.63 .87 B 93 11/5 5 Wet 33 89 611 447 6.55 .47 B 117 11/10 5 Dry V 59 86 571 418 5.30 1.20 C 137 12/8 6 Wet 25 76 - C 150 12/15 7 Wet 14 42 B 72 11/2 8 Wet 92 90 653 437 5.49 1.26 B 114 11/10 8 Dry / 209 89 644 483 5.51 1.14 B 113 11/10 9 Wet V 21 79 546 352 6.35 1.80 Continued on next page Appendix Table 7 Continued. Collection Percent Fat Body Weights Total Ovarian/ Kidney Metatarsal Weight of cg Adrenal per Area Date Age Uterine OUndSi Fat Marrow Adrenals lb Hog-dressed 68- No. (1968) (Years) Udder Data Index (Wet Basis) Whole Hog-dressed (g) Body Weight B 123 11/23 11-15 Wet 12 14 551 417 6.68 1.60 B 124 12/14 11-15 Wet 23 77 605 377 4.70 1.27 C 132 12/7 11-15 Wet J,J 7 57 -

A North Coast area B = Millicosna area C = Loon Lale area 'Corpusluteuin present. 'Uteruscontaining nonviable chorion. Ovaries containing cystic follicles.